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BULLETIN OF 

THE BRITISH MUSEUM 

(NATURAL HISTORY) 



ZOOLOGY 

VOL. I 

1950-1953 



PRINTED BY ORDER OF THE TRUSTEES OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

LONDON : 1954 



DATES OF PUBLICATION OF THE PARTS 



No. 


i. 


17 January 1950 


No. 


2. 


20 March 1950 


No. 


3- 


31 March 1950 


No. 


4- 


29 August 1950 


No. 


5- 


3 November 1951 


No. 


6. 


26 October 195 1 


No. 


7- 


6 June 1952 


No. 


8. 


5 August 1952 


No. 


9- 


31 July 1952 


No. 


10. 


11 December 1952 


No. 


11. 


13 January 1953 


No. 


12. 


27 November 1953 







PRINTED IN 

GREAT BRITAIN 

AT THE 

BARTHOLOMEW PRESS 

DORKING 

BY 

ADLARD AND SON, LTD. 



CONTENTS 

ZOOLOGY VOLUME I 

No. I. On some species of Lernaea (Crustacea, Copepoda : parasites of fresh- 
water fish). By J. P. HARDING I 

No. 2. On a giant squid, Ommastrephes caroli Furtado, stranded at Looe, 

Cornwall. By w. j. rees (Pis. 1-2) 29 

No. 3. The identity of Captain Cook's kangaroo. By t. c. s. morrison- 

scott and f. c. sawyer (Pis. 3-5) 43 

No. 4. Notes on Asteroids in the British Museum (Natural History). By 

D. DILWYN JOHN (PL 6) 51 

Lernaeodiscus pusillus nov. spec, a Rhizocephalan parasite of a Pov- 
cellana from Egypt. By hilbrand boschma 6i 

No. 5. On a rare deep-sea fish Notacanthus phasganorus Goode (Heteromi- 
Notacanthidae) from the Arctic Bear Isle fishing grounds. By d. w. 
tucker and J. w. jones (Pis. 7-9) 67 

No. 6. The ocean sunfishes (family Molidae) . By a. fraser-brunner 8y 

No. 7. The cestodes of seals from the Antarctic. By stanislaw markowski 

(Pis. 10-21) 123 

No. 8. The ' Manihine' Expedition to the Gulf of Aqaba 1948-1949 

Foreword . Station list and collectors' notes (Pis. 22-27) J 53 

Preliminary hydrological report. By G. e. r. deacon 159 

Sponges. By maurice burton 163 

Turbellaria . Polycladida. By Stephen prudhoe 175 

Gephyrea. By A. c. Stephen 181 

Mollusca. By w. j. rees and A. stuckey (Pis. 28-30) 183 

Echinodermata. By ailsa m. clark (Pis. 31-32) 203 

Tunicata. By willard g. van name 215 

Fishes. By n. b. Marshall 221 

No. 9. On the species and races of the yellow wagtails from Western Europe 
to Western North America. By c. h. b. grant and c. w. mackworth- 

PRAED (Pis. 33-35) 253 

No. 10. Mammals collected by Mr. Shaw Mayer in New Guinea 1932-1949. 

By ELEANOR M. O. LAURIE 269 

No. 11. Taxonomy of the karroo and red-back larks of Western South Africa. 

By J. D. MACDONALD (Pis. 36-38) 319 

No. 12. Suberites domuncula (Olivi) : its synonymy, distribution, and ecology. 

By m. burton 353 

Notes on Asteroids in the British Museum (Natural History) III & IV. 

By A. M. CLARK (Pis. 39-46) 379 

Some inter-tidal mites from south-west England. By g. o. evans 
and E. browning 413 

Index to Volume I 423 



ERRATUM 
Plates 7-9. 
For NOTOCANTHUS read NOTACANTHUS. 



.SSliSlW .1 7 JAN iS5U 

ON SOME sl*£OES OF 
LERNAEA 

(CRUSTACEA, COPEPODA: 
PARASITES OF FRESH-WATER FISH) 

J. P. HARDING 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ZOOLOGY Vol. i No. i 

LONDON: 1950 



ON SOME SPECIES OF LERNAEA 

(CRUSTACEA, COPEPODAI 
PARASITES OF FRESHWATER FISH) 



BY 

J. P. HARDI NG 



% 




Pp. 1-27; gs Text-figures 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) y 
ZOOLOGY Vol. 1 No. 1 

LONDON: 1950 



THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted in 1949, is to be 
issued in five series, corresponding to the Departments 
of the Museum. 

Parts will appear at irregular intervals as they be- 
come ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

This paper is Vol. 1, No. 1, of the Zoological series. 



PRINTED BY ORDER OF THE TRUSTEES OF 
THE BRITISH MUSEUM 

Issued January 1950 Price Seven shillings and sixpence 



ON SOME SPECIES OF LERNAEA (CRUSTACEA, 
COPEPODA: PARASITES OF FRESHWATER FISH) 

By J. P. HARDING 

(With ninety-five text-figures) 

SYNOPSIS 

Twenty-eight species of Lernaea are recognized, of which fourteen are represented in the collections of 
the British Museum. Nine of these are new species. In addition there are seven names in the literature 
which are relegated to the synonymy. A key to the twenty-eight recognized species is given and the 
fourteen species in the Museum are described and figured. 

One result of the renewed interest in the freshwater fisheries of Africa and other 
countries in recent years has been an accumulation of the parasites of these fish in 
the British Museum with requests for their identification. Attempts to name the 
species of Lernaea on the basis of existing descriptions and keys soon showed that 
a high proportion of them were new, and that Cunnington's (1914) and Wilson's 
(1917 & 1918) revisions and keys are out of date. 

There is no need here to repeat recriminations against Wilson (1917) for transposing 
the names for the genera Lernaea and Lernaeocera. The inevitable confusion caused 
by this strict interpretation of the Rules of Nomenclature has fortunately been 
lessened by the fact that subsequent workers have, however unwillingly, nearly 
all agreed to follow. 

As is often the case with degenerate parasitic forms, the characters used to distin- 
guish between species are often ill-defined and not easily seen. Wilson often expressed 
the opinion that species of Lernaea and other parasitic copepods could readily be 
distinguished by reference to their appendages; but I have found extraordinarily 
little difference between the appendages of one species and those of another. The 
first four pairs of pereiopods, which will be referred to as legs 1 to 4 in this paper, 
are the only appendages that are easily examined as they are flat, and as each has a 
number of setae and spines I expected variations in their arrangements to provide 
useful characters for distinguishing species. Unfortunately I could find hardly any 
variations. The precise arrangement of the setae and spines was investigated in 
fourteen species, and in thirteen of them it was identical, only one of them could be 
separated on this basis. Table 1 gives the arrangement of the setae and spines in 
the thirteen species. L. bistricornis is included although I was able to see only legs 3 
and 4; all four pairs of legs were seen in the other species. L. haplocephala differs 
from all these species in having four setae instead of five on the terminal segment 
of each exopod (Table 2, p. 19). L. oryzophila, according to Monod's (1932) descrip- 
tion and figures, differs from the other species, having four instead of three spines 
on the last segment of the second exopod and two spines instead of three on the 
last segment of the fourth exopod. L. dolabroides (Wilson, 1918, figs, jy and 78) 
seems to have a quite different setation of its legs. 



4 ON SOME SPECIES OF LERNAEA 

Table i 

Arrangement of the setae and spines on the legs of L. bagri, L. barbicola, L. barilii, 

L. barnimiana, L. bistricornis, L. cyprinacea, L. dicer acephala, L. longa, L. lophiara, 

L. palatae, L. piscinae, L. tilapiae, and L. tuberosa 



Exopod {^f 
End °P° d { setae' 



Leg i 



1. 1. 2 

1. 1. 5 

0.0.2 

1. 1.4 



Leg 2 



1. 1. 3 
1. 1.5 

0.0.2 
I.2. 4 



Leg 3 



1. 1. 3 

1. 1. 5 

0.0.2 
I.2. 4 



Leg 4 



1. 1. 3 
1. 1-5 

0.0.2 

1.2.3 



The other appendages are even less useful than the legs for separating one species 
of Lernaea from another. Wilson (1920, p. 7) claims that L. haplocephala may be 
distinguished by the 'small spherical terminal joint of the maxillipedes, with its four 
curved claws \ This may have been true of the specimen he examined, but I find that 
the maxillipedes of the type specimens of L. haplocephala have five claws like any 
other species and that the terminal segment does not appear to have any specific 
shape. Fig. 34 gives the arrangement of the mouth parts as far as I have been able 
to see these very minute appendages. 

We are left with the shape of the body and with the internal anatomy for distin- 
guishing species. Unfortunately it is difficult to study one without destroying, or 
at least distorting, the other, and I have neglected a study of the internal anatomy 
in favour of the shape of the body and its processes, and in particular that of the 
anchor. I am restricting the use of the word 'head' to that small, rounded part 
which bears the antennae and the mouth parts. The swollen part with processes 
which are usually described as 'cephalic processes', 'cephalic arms', or 'cephalic 
horns ' I propose to call the anchor and its arms, as this describes both the appearance 
and the function of this part of the body. The anchor is often difficult to remove 
from the flesh of the fish without damage, and I have adopted the method of cutting 
out the part of the fish with the parasite embedded in it and placing it in a tube 
with a solution of potassium hydroxide to which a little chlorazol black has been 
added. If this is left for about twenty-four hours the tissues of the fish are usually 
softened sufficiently for the Lernaea easily to be removed ; the chlorazol black stains 
the chitin of the parasite a dark blue. The external shape of the animal is well 
preserved by this method and the appendages can be examined without difficulty. 
The egg-sacs should be removed before placing the parasite in the hydroxide or they 
will be destroyed. 

All drawings and measurements of specimens recorded in this paper have been 
made with the aid of a camera lucida. The total length of a specimen is understood 
to mean the length from the front of the head to the end of the abdomen, allowance 
being made for bends and curves in the body. Parts of the anchor which may project 
in front of the head and the f ureal setae are not included in this measurement. The 
measurement and drawing of curved specimens was helped by the use of gimbals 
which enabled the specimen to be held in any position on the stage of the microscope. 
The gimbals, similar in principle to those of a ship's compass, but with sufficient 



ON SOME SPECIES OF LERNAEA 5 

friction in the bearings to prevent free swinging, were made of concentric cylinders 
of perspex, as shown in Fig. I. The whole is submerged in formalin in a glass vessel 
and the specimen is placed in the central cylinder, which is closed at the bottom 
to form a dish. 

I have found the shape of the anchor and its arms to provide the most useful 
characters for taxonomic purposes. In spite of the fact that the shape of the anchor 
is liable to be distorted by meeting bones and other hard obstructions during its 
growth in the body of the fish, each species has a characteristic form which varies 
within limits which are usually definable provided sufficient material is available. 




Fig. 



i. Gimbals for tilting the specimen into positions required for 
drawing or measuring. 



The abdomen and the pregenital prominences of each species examined have been 
drawn with some care from more than one aspect as I have found them useful in 
separating species. The shape of these parts is not so easily influenced by the site 
of attachment of the parasite, but on the other hand there is often a difference in 
the shape of these parts of a young individual and those of an old one. The characters 
of the immature specimens are often less well defined than those of adult specimens, 
and the adult female seems to continue to grow and develop specific form for some 
time after eggs are first produced. 

The positions of the legs on the body have been recorded wherever possible with 
an estimate of the range of variability. 

The amount of torsion and its direction is very variable; but worth recording 
because some species show little torsion and in others considerable torsion is the rule. 
In some species, such as L. barnimiana, the torsion is not only variable in direction 
and extent in different specimens but often changes its direction along the length 
of the one individual. 

Cunnington (1914) remarks on the rarity of copepod parasites on the fish he 
collected from Lake Tanganyika. Lake Nyasa, however, seems to be very rich in 
species. More information is required about the seasonal distribution of the different 
species. Miss R. H. Low tells me that she found Lernaea on many of the specimens of 
Bagrus that she collected in August and that by November the Bagrus were free of 
parasites but the Tilapia were infected. She had the impression that the parasites 



6 ON SOME SPECIES OF LERNAEA 

had transferred their attentions from Bagrus to Tilapia ; but an examination of the 
specimens shows that there are two distinct species of Lernaea involved, L. bagri 
and L. tilapiae, described below. 

Lernaea cyprinacea Linnaeus 
Figs. 2-12. 

1746 Levnea tentaculis quatuor Linnaeus, Fauna Svecica: 367, pi. 2. 

1758 Lernaea cyprinacea Linnaeus, Syst. Nat.: 655. 

1783 ,, ,, : Barbut, Gen. Vermium: 67, pi. 7, fig. 3. 

1822 Lemeocera cyprinacea (Linnaeus) Blainville, Journ. Phys. 95: 377. 

1835 Lernaeocera ,, : Burmeister, Nova Acta Leop. Carol. 17: 309, pi. 24 a, figs. 1-3. 

1850 ,, ,, : Baird, Brit. Entomost: 343, pi. 35, fig. 13. 

1904 ,, ,, : Hofer, Handb. Fischkrankheit, Munchen: 144, fig. 95 and p. 119 

[fide Pesta 1934]. 
1909 ,, ,, : Neresheimer, Brauer: Susswasserfauna Deutschl. 11: 77, fig. 326. 

1913 ,, ,, (part): T. and A. Scott, Brit. Parasit. Cop., Ray Soc. London: 

154, pi. 50, figs. 4-5 [not figs. 1-3]. 

1917 Lernaea cyprinacea: Wilson, Proc. U.S. Nat. Mus. 53: 4, 39. 

1918 ,, ,, : Wilson, Bull. U.S. Bur. Fish. 35: 193, 196, pi. 15, fig. 86. 
1925 ,, (Lernaeocera) elegans: Leigh-Sharpe, Parasitology, 17: 245, text-figs. 1-5. 
1927 ,, elegans: Nakai, /. Fish. Inst. Tokyo, 23: 39, pis. 2-4, text-figs. 1-7. 

1927 ,, cyprinacea: Okada, Annot. Zool. Jap. Tokyo, 11: 185, text-figs. 1-2. 

1928 ,, elegans'. Matsui and Kumada, /. Fish. Inst. Tokyo, 23: 101, pis. 5-7. 

1932 ,, cyprinacea: Monod. Ann. Parasit. hum. comp. 10: 362, text-figs. 8 H, 11, 12. 

1933 ,, ,, : Gurney, Brit. Fresh-water Cop, Ray Soc. Lond. 3: 338 ,text-figs. 1969, 

1971-1983. 

1933 ,, carassii Tidd, Ohio J. Sci. 33: 465, pi., figs. 1-8. 

!934 ,> cyprinacea: Markewitsch, Ann. Mus. zool. polon. 10: 234, pi. 45, fig. 8. 

1934 >> >> ' Pesta, Tierwelt Deutschl. 29: 42, text-fig. 25. 

1937 y> >> '• Markewitsch, Cop. Parasit. Binnengewdss. U.S.S.R., Kiev: 98, pi. 8. 

1937 >> >> : Wagler, Tierwelt Mitteleuropas, Crust. 2, 2a: 179, text-fig. 542. 

1939 ,, ,, : Yamaguti, Vol. Jubil. Prof. S. Yoshida 2: 475, pi. 30, figs. 156-165. 

The material in the British Museum consists of two specimens found by Dr. Gurney 
on a specimen of Carassius carassius (L.) from Sweden in the Museum fish collection ; 
a specimen from Canon Norman's collection which was labelled ' L. esoscina from 
Prof. Heller ' (this specimen is unfortunately without record of host or locality) ; 
thirty or more specimens from Japan presented by Dr. Gurney, and finally a few 
microscope slides of the type specimens of L. elegans presented by Mr. Leigh-Sharpe. 

I have little to add to the excellent descriptions and figures given by many of the 
authors listed above, Gurney (1933) in particular ; the shape of the anchor is, however, 
rather more variable than these descriptions indicate. The most typical arrangement 
is that of the Swedish specimen (Fig. 2), if the right dorsal arm which is distorted is 
ignored ; the arms are all rather long and slender and the dorsal arms are T-shaped. 
This is the arrangement shown in nearly all figures of European specimens from 
Linnaeus, 1746 to Monod, 1932, and Gurney, 1933. Very few of the Japanese speci- 
mens are quite like this, there is a tendency for the dorsal arms to be Y-shaped 
(Figs. 5-7), and the posterior fork of the Y is often reduced. Prof. Heller's specimen 
(Figs. 10-12) of unknown origin is very like the Japanese specimens . The pregenital 



ON SOME SPECIES OF LERNAEA 



prominence of L. cyprinacea is generally described as 'simple or only slightly in- 
dented' (Wilson, 1918, p. 193, key). It is simple in the Swedish specimens (Figs. 3 
and 4). The Japanese specimens have a distinctly double pregenital prominence 
(Figs. 8 and 9). Prof. Heller's specimen again agrees with the Japanese specimens 




Figs. 2-12. Lernaea cyprinacea Linnaeus 

Figs. 2-4, specimen from Sweden; Fig. 2, dorsal view of anchor; Fig. 3, ventral view of pregenital prominence 
and abdomen; Fig. 4, lateral view of the same; Figs. 5-9, specimens from Japan; Figs. 10-12, specimen from 
Prof. Heller. The line near each figure is 1 mm. drawn to the same scale. 

rather than with the Swedish ones. Markewitsch (1937) also gives a figure of a speci- 
men with a bilobed pregenital prominence, but he does not say from what part of the 
U.S.S.R. it came. None of the characters of L. elegans given by Leigh-Sharpe are 
valid for distinguishing between the Japanese form and the European. No ' auricular 
expansions' are now visible on the specimen from which Leigh-Sharpe 's Fig. 3 was 
based ; possibly the artist saw some folds of chitin. In this figure are shown what are 
called 3-segmented uncinate thoracic appendages, but an examination of the speci- 
men shows that these are not appendages at all but are the badly fixed internal 
tissues which can be seen only by focusing well below the surface of the body, as 
Monod (1932) has already pointed out. Leigh-Sharpe's types of L. elegans are the 
same form of L. cyprinacea as the Japanese specimens I have seen. 



8 ON SOME SPECIES OF LERNAEA 

I have looked at the appendages of the Swedish and Japanese specimens with 
some care, but have been unable to find any difference, the setation of the legs is 
precisely the same in both (Table i). The positions of the legs on the body of five 
Japanese specimens were measured; there was considerable variation, particularly 
of the anterior legs, but the positions of the five pairs of legs do not enable the 
Japanese and European specimens to be separated from one another. The positions 
of the five legs were 6-9, 16-20, 42-45, 73-74, and 90-92 per cent, of the total body 
length distant from the most anterior part of the head, respectively. 

There is little doubt that L. carassii Tidd is the Japanese or elegans form of 
L. cyprinacea. 

Lernaea barnimiana (Hartmann) 
Figs. 13-28 

1865 Lernaeocera barnimiana Hartmann, Naturges.-med. Skizze Nillander: 206. 

1870 ,, barnimii Hartmann, Arch. Anat. Phys. Wiss. Med. 1870: 726, pis. 17-18. 

1871 ,, ,, : Hartmann, 5. B. naturf. Fr. Berl.: 60. 

1914 ,, temnocephala Cunnington, Proc. Zool. Soc. Lond. 1914: 827, pi. 1, figs. 8-9, 

text-fig. 1 c. 

1917 Lernaea barnimii: Wilson, Proc. U.S. Nat. Mus. 53: 38. 

1918 ,, temnocephala: Wilson, Bull. U.S. Bur. Fish. 35: 193, 196, pi. 15, fig. 87. 
1918 ,, barnimii: ibid.: 193, 196, pi. 15, fig. 94. 

1940 ,, temnocephala: Brian, Boll. Idrobiol. Caccia Pesca, 1: 50, pi., figs. A-F. 

1944 ,, barnimiana: Capart, Bull. Mus. Hist. nat. Belg. 20 (24): 2, text-fig. 1. 

Several specimens of this species were taken from a fish, Labeo forskalii Ruppell, 
caught in Lake Edward by Dr. E. B. Worthington in 193 1. The heads of the parasites 
were buried in the flesh of the head and inside the mouth of the fish. The Museum 
also possesses the single specimen on which Cunnington founded L. temnocephala. 
Thanks to the kindness of Dr. Capart I have seen three of the specimens he described 
from the Belgian Congo. 

The length of the adult female, judging from the literature, ranges from 7 mm. to 
12 mm. (Capart's 1944 figs.) ; Hartmann's 1870 record of a range of from 10 mm. to 
14 mm. may include the anterior arms of the anchor in the length. The British 
Museum specimens range from 8-2 mm. to io-8 mm. in length. 

The positions of the five pairs of legs of seven of the Lake Edward specimens give 
the following ranges measured in percentages of the total body length: 7-2-9-8, 
19-24, 41-51, 73-79, and 89-92 per cent, respectively. The positions of the legs on 
the specimens kindly lent me by Dr. Capart agree, but two other specimens which 
he figures (Capart, 1944, fig. 1, A and e) appear to have the first and second legs a 
little farther forward ; this may, however, be owing to the foreshortening which is 
inevitable when curved specimens are drawn. Hartmann's (1870) drawings are not 
very reliable and I attach no importance to the fact that the position of the fourth 
pair of legs in his Fig. 1 is only 65 per cent, of the body length from the anterior. 
The positions for Cunnington 's type of L. temnocephala are 8, 18, 44, 79, and 92 
per cent, respectively. The variations in the positions of the legs seems to be quite 
independent of the size of the specimen, i.e. there is no heterogony with respect to 
this character. 



ON SOME SPECIES OF LERNAEA 



The torsion of the specimens I have seen was variable ; that between successive 
pairs of legs never exceeded 90 ° and was usually much less. It could be either sinistral 
or dextral and frequently changed its direction. The total torsion of the whole body 
was not more than 120 in any of the twelve specimens examined. 




Figs. 13-28. Lernaea barnimiana (Hartmann). 
Figs. 13-16, Cunnington's specimen from L. Tanganyika, named by him temnocephala ; Fig. 13, ventral view of 
anchor; Fig. 14, dorsal view; Fig 15, ventral view of abdomen and pregenital prominence; Fig. 16, lateral view 
of the same. Figs. 17-28, specimens from L. Edward. Fig. 17, ventral view of anchor; Fig. 18, ventral view of 
another specimen; Fig. 19, anterior view of the latter; Figs. 20 and 21, posterior end of this specimen; Fig. 22, 
posterior end of a specimen with egg-sacs; Figs. 23-28, anchors of other specimens from L. Edward, all from the 
same fish. The line near each figure is 1 mm. drawn to the same scale. 

The arms of the anchor are rather variable in shape and arrangement, as Capart 
(1944) has shown. The most usual arrangement is for the part between the head and 
the first legs to be swollen and more or less globular. The ventral arms are simple in 
shape and very short. The usual arrangement is for the ventral arms to be directed 
outwards ; this was so in all the Lake Edward material, in Cunnington's specimen 
from the Nile, and in most of Capart's material from the Belgian Congo. In some 
of Capart's specimens and also in the figure accompanying Hartmann's description 
(Hartmann, 1870, fig. 1), on the other hand, the ventral processes are directed ante- 
riorly. With regard to the bifurcating dorsal arms, Hartmann's figure shows both 
branches equal to one another and both diverging slightly away from the body; 
but in his description he says that the anterior branch is the longer of the two and 

zoo. 1, 1. B 



io ON SOME SPECIES OF LERNAEA 

is the more outwardly directed. The normal condition seems to be for the anterior 
branch to diverge widely from the body while the posterior branch is directed slightly 
inwards (Figs. 17, 18, 23, &c). The angle between the two branches is normally an 
open and continuous curve ; but sometimes as in Figs. 25, 27, 28, &c. there is a more 
or less distinct angle. The Y-shaped condition of the arms of the temnocephala holo- 
type (Fig. 14) is unusual but within the range of variation of L. barnimiana. 

The pregenital prominence is distinct and bilobed (Figs. 15, 16, 20, and 21) ; but 
from some aspects it may appear to be a single broad process. 

The abdomen is distinctly 3-segmented and may continue the line of the body 
or be set at an angle. Each segment is a little smaller than the preceding one. 
Hartmann does not describe the abdomen of his specimens, and his figure and those 
of Brian (1940) are of little value in this respect. Cunnington's temnocephala specimen 
(Figs. 15 and 16) is normal. 

The setation of the legs is the same as that of L. cyprinacea (Table 1, p. 4). I have 
cleared Cunnington's type of temnocephala with potassium hydroxide, and this has 
got rid of the twists and distortions he mentions and has enabled me to examine the 
setation of its legs; and as with the other characters investigated I can find no 
difference between this specimen and the Lake Edward specimens and I have no 
hesitation in placing L. temnocephala (Cunnington) in the synonymy of L. barnimiana, 
as Capart (1944) has already suggested. 

Lernaea piscinae sp. nov. 
Figs. 29-34 

Holotype, Reg. No. 1949.8.14.1, and many paratypes, all females, in the British 
Museum. The parasites were found heavily infesting a Cyprinid fish, Hypophthal- 
micthys nobilis (Richardson) cultivated by the Chinese on a fish farm at Singapore. 
Four fish heavily infested with the parasites, over 50 per fish, were presented to the 
Museum by Mr. W. Birthwhistle in 1929. Length of holotype 10*4 mm. ; the length 
of 10 paratypes ranged from 97 mm. to 12-4 mm. The positions of the five pairs of 
legs of these eleven specimens were 5-7, 13-14, 31-38, 69-74, and 91-93 per cent, 
of the total length from the most anterior part of the head. All the specimens of this 
species were very much alike ; seven out of the eleven specimens had a curve between 
legs 2 and 3 as in Fig. 29. Three of the remainder were straight and the other had an 
additional bend between legs 1 and 2. 

The main part of the anchor forms a bar set at right angles to the body like the 
cross-bar of a T (Figs. 29 and 30). The middle of the bar is considerably thicker than 
the part of the body joined to it, and tapers gradually towards the ends, which are 
also curved slightly in an ant ero- ventral direction. From about the middle of each 
half of the cross-bar there is a short dorsal process. There is also a pair of ventral 
processes between the head and the legs 1 ; these are separated by a distance about 
equal to the width of the head and are directed slightly inwards towards one another. 

Except for slight swellings at the positions of the legs the body increases in thick- 
ness very gradually from before backwards. 

The abdomen (Figs. 32-33) makes a slight angle with the body ; it is nearly 1 mm. 



ON SOME SPECIES OF LERNAEA 



ii 



long and less than J mm. wide. Ventrally it is distinctly 3-segmented ; but the dorsal 
profile forms an even, continuous curve. 

The pregenital process is double, the two lobes being small but well defined and 
quite separate from one another. 




Figs. 29-34. Lernaea piscinae sp. nov. 

Fig. 29, ventral view of holotype ; Fig. 30, anterior view of anchor ; Fig. 31, lateral view of posterior end of a paratype 

with egg-sacs; Fig. 32, ventral view of abdomen and pregenital prominences of holotype; Fig. 33, lateral view of 

the same ; Fig. 34, mouth and associated appendages. l.L, lower lip ; md., mandible ; mxa., maxilla ; mxl., maxillule. 

The line near each figure is 1 mm. drawn to the same scale except that near fig. 34 which is 0-02 mm. 

The egg-sacs (Fig. 31) are very long, about 4 mm., three-quarters of their length 
projecting beyond the tip of the abdomen. 

The setation of the legs is the same as in L. cyprinacea (Table 1). 

The mouth parts (Fig. 34), as far as I was able to make out, are the same as for 
other species. 

Lernaea diceracephala (Cunnington) 
Figs. 35-39 

1914 Lernaeocera diceracephala Cunnington, Proc. Zool. Soc. Lond. 1914: 824, pi. 1, figs. 1-3, 
text-fig. 1 a. 

1917 Lernaea diceracephala: Wilson, Proc. U.S. Nat. Mus. 53: 38. 

1918 ,, ,, : Wilson, Bull. U.S. Bur. Fish. 35: 192, 194, pi. 15, fig. 90. 
1944 ,, ,, : Capart, Bull. Mus. Hist. nat. Belg. 20 (24): 7. 

Holotype, Reg. No. 1914.12.2.1, and one paratype in the British Museum; I have 



12 ON SOME SPECIES OF LERNAEA 

selected the more perfect of the two specimens as the holotype. These are the only 
specimens known and were taken from the gill arches of a large Clarias mossambicus 
Peters, caught at Sumbu, Lake Tanganyika, by Dr. Cunnington in 1904. Capart 
(1944) includes the species in his paper because part of Lake Tanganyika lies in the 
Belgian Congo. 

Cunnington's description of the two specimens is very good ; but he describes the 
left arm as being complete in the better specimen when in fact the tip has been 
broken off. 

The length of the holotype measured as if straightened out is 9-1 mm. The five 
pairs of legs come in positions 10, 23, 50, 71, and 92 per cent, of the total length from 
the most anterior part of the head. 

I have made drawings of the two specimens which I hope are an improvement on 
Cunnington's photographs, and which show how similar to one another are the bends 
and constrictions in the body. 

Lernaea bagri sp. nov. 
Figs. 40-43 

Holotype, Reg. No. 1949.8. 14. 9, and over two dozen paratypes, all females, in the 
British Museum. 

The copepods were taken from Bagrus meridionalis in Lake Nyasa by Miss R. H. 
Low, 14 Aug. and 22 Sept. 1946. 

The length of the holotype is 12 -i mm. ; that of twenty-four adult females carrying 
egg-sacs ranged from 9-9 mm. to 14-2 mm. 

The body of a few of the slender, young-looking specimens is straight, but usually 
it is curved as shown (Fig. 40), and in these there is a torsion which in this species 
nearly always changes its direction. In all the specimens I have examined for this 
purpose the torsion is first sinistral and then dextral. In the holotype, for example, 
the torsion between legs 1 and 2 is 40 sinistral, between legs 2 and 3 it is 8o° dextral, 
and between legs 3 and 4 it is a further 50 dextral, after which there is no further 
torsion. The resultant torsion between the head and the abdomen is about 90 dextral. 

The arms of the anchor are heavily chitinized, in contrast to those of the next 
species to be described, L. tilapiae, and lie in a plane approximately at right angles 
to the body. The head is placed centrally over the cross formed by the four arms. 
The ventral arms are straight and the dorsal ones are curved towards them. There is 
a tendency for each arm to end with a rounded knob. 

The positions of the legs are rather variable in this species. Six specimens were 
examined, and the positions of legs 1 to 5 gave the following ranges respectively: 
7-10, 18-22, 42-52, 73-78, and 90-93 per cent. The setation of the legs is the same as 
that of L. cyprinacea (Table 1). 

The abdomen (Figs. 41-3) is set at a slight angle to the line of the body ; it is 
straight and slightly tapering ; the three segments are very indistinctly separated. 

The pregenital prominence is bilobed. Sometimes the lobes are prominent and 
bulge laterally beyond the greatest width of the body, but usually, as in the holotype, 
they are not very prominent from the ventral aspect (Fig. 43). 



ON SOME SPECIES OF LERNAEA 



13 



The egg-sacs (Fig. 41) are about z\ mm. long and \ mm. wide at their greatest 
width, which lies at about the proximal third of the length. 




Figs. 35-39. Lernaea diceracephala (Cunnington) . 

Fig. 35 > ventral view of holotype; Fig. 36, dorsal view of abdomen; Fig. 37, constriction between legs 3 and 4 
from a slightly different aspect from that of Fig. 35 ; Fig. 38, lateral view of paratype; Fig. 39, similar view of 

part of holotype. 

Figs. 40-43. Lernaea bagri, sp. nov. 

Fig. 40, dorsal view of holotype; Fig. 41, lateral view of abdomen and egg-sacs of a paratype; Fig. 42, lateral view 
of abdomen and pregenital prominences; Fig. 43, ventral view of the same. The line near each figure is 1 mm. 

drawn to the same scale. 



Lernaea tilapiae sp. nov. 
Figs. 44-46 

Holotype, Reg. No. 1949. 8. 14.17, and a few paratypes, all females, in the British 
Museum. 

The parasites were collected by Miss R. H. Low from Lake Nyasa and were taken 
from the mouth and gills of Tilapia squamipinnis Giinther and T. lidole Trewavas 
caught in Lake Nyasa 22 Nov. 1946. 

The length of the holotype measured from the front of the head to the tip of the 



M 



ON SOME SPECIES OF LERNAEA 



abdomen is 9-2 mm. Five other females bearing egg-sacs ranged from 7-5 mm. to 
11 mm. in length. 

The body is comparatively slender from the head to as far as legs 3 and is usually 
curved here, so that the anterior part of the body is at right angles to the broad part 
behind legs 3 (Fig. 44). In the holotype the torsion is dextral 45 ° between legs 2 and 3, 




Figs. 44-46. Lernaea tilapiae sp. nov. 

Fig. 44, dorsal view of holotype; Fig. 45, lateral view of abdomen, pregenital prominence, and egg-sacs; Fig. 46, 
ventral view of the same without egg-sacs. The line near each figure is 1 mm. drawn to the same scale. 

dextral 90 between legs 3 and 4, and dextral a few degrees beyond legs 4, the total 
torsion being dextral through about 140 . The only torsion in one of the paratypes 
is a sinistral one of 45 ° between legs 3 and 4. 

The anchor bears four long straight slender arms as figured (Fig. 44) ; these lie in 
a plane nearly parallel to that of the body ; the posterior pair are directed backwards 
and are only slightly divergent; the anterior pair diverge widely, with the head 
placed in the angle between them. The four arms of the anchor are about equal in 
length to one another and more than half the length of the body. They are only 
lightly chitinized and are much softer than those of the last species described, L. bagri. 



ON SOME SPECIES OF LERNAEA 15 

The legs come in positions 8, 25, 50, 77, and 90 per cent, of the body length from 
the anterior end. The setation of legs 1 to 4 is the same as for L. cyprinacea (Table 1, 

P-4)- 

The abdomen is divided into three segments by transverse ventral constrictions 

which give it a characteristic profile (Fig. 45). The dorsal profile of the abdomen is 

slightly arched. 

The pregenital prominence is bilobed; the two lobes overhang the abdomen 
slightly, but their ventral surface is in line with that of the body in front (Fig. 45). 

The egg-sacs are about 2-5 mm. long and 0-5 mm. wide, slightly tapering towards 
each end. Miss Low records that in life the parasite is brown in colour and the eggs 
are jade-green. 



Lernaea barilii sp. nov. 
Figs. 47-60 

Holotype, Reg. No. 1949.8. 14.21, and about 10 paratypes, all females, in the 
British Museum. 

The parasites were taken on a large specimen (500 mm. long) of Barilius microlepis 
Giinther from Lake Nyasa by Dr. Christy in 1925, a piece of the flank of the fish 
with the copepods embedded being preserved together with a note to the effect that 
there were more parasites on the tongue, &c. I have only seen the specimens from 
the flank. 

The length of the holotype is 8-3 mm. with the positions of legs 1 to 5 at 8-4, 20, 
47, yy, and 92 per cent, of the total body length from the anterior end respectively. 
The positions on paratype Reg. No. 1949.8. 14.24 are 8, 19, 47, yy, and 93 per cent. 
The setation of the legs is the same as in L. cyprinacea (Table 1, p. 4). 

The body is straight and short, widest at the posterior end. In the two specimens 
which were examined in detail the torsion was about 80 °. In the holotype most of 
the torsion was between legs 2 and 3, and in the paratype examined it was between 
legs 3 and 4 ; it was sinistral in the holotype and dextral in the paratype. 

The arrangement of the anchor is best understood with reference to Figs. 47-49 
and 53-55. There are a pair of lateral T-shaped arms with the cross-bar of the T 
running more or less parallel to the body ; the basal part of these arms is short and 
thick. Anterior and ventral to the dorsal arms are a pair of simple arms directed 
outwards, with in nearly all cases a small knob facing anteriorly. 

The pregenital prominences are very large and distinctly separated from one 
another; they reach almost to the end of the abdomen in some specimens (Figs. 40, 
50-52, 57-6o). 

The abdomen, particularly the part composed of the last two segments, is very 
small and set at an angle to the body. All three segments are clearly marked off 
from one another ventrally; the first is very much broader than the other two 
(Figs. 51 and 60). 

The egg-sacs of the holotype were about 2.75 mm. long, broadest in the middle 
and tapering towards each end (Fig. 58). 



i6 



ON SOME SPECIES OF LERNAEA 




Figs. 47-60. Lernaea barilii sp. nov. 
Fig. 47, ventral view of holotype; Fig. 48, dorsolateral view of anchor; Fig. 49, lateral view of the same; Fig. 50, 
dorsal view of posterior end of holotype; Fig. 51, lateral view of the same; Fig. 52, ventral view of the same; 
Fig. 53, ventral view of anchor of a paratype; Fig. 54, anterior view of another paratype; Fig. 55, ventral view 
of the same; Fig. 56, anchor of a specimen drawn in situ by clearing in benzyl alcohol; Fig. 57, lateral view of 
specimen with egg-sacs before removing from the fish (the specimen has shrunk and collapsed dorsally); Fig. 58, 
ventral view of holotype with egg-sacs before treatment with hydroxide; Fig. 59, ventral view of the posterior 
end of a paratype; Fig. 60, lateral view of the same. The line near each figure is 1 mm. drawn to the same scale. 

Lernaea palati sp. nov. 
Figs. 61-64 

Holotype, Reg. No. 1949.8. 14.26 in the British Museum. The single specimen on 
which this species is based was from the roof of the mouth of a fish, Haplochromis 
chrysonotus (Boulenger) from Vua on Lake Nyasa, collected by Dr. Christy in 1925. 
The hind end of the parasite projected through a gill slit and was visible externally. 

The length of the specimen, allowance being made for bends, is 127 mm. The 
body from the first pair of legs to half-way between legs 2 and 3 is cylindrical, 
about 07 mm. thick ; the section containing legs 3 is broader, about 1*2 mm. across ; 
there is a waist between legs 3 and 4 ; the body bends backwards and to the left here 
and bulges again to a thickness of 1-2 mm. in front of legs 4. 

The abdomen is tilted dorsally at an angle of about 45 ° ; it is a simple cylinder 
rounded at the end about 1 mm. long and 07 mm. broad without any sign of seg- 
mentation. 



ON SOME SPECIES OF LERNAEA 



17 



The five pairs of legs are placed in positions 87, 26, 55, 80, and 93 per cent, of 
the body length from the anterior end. The setation is the same as that of L. cypri- 
nacea (Table 1, p. 4). There is little torsion. 




Figs. 61-64. Lernaea palati sp. nov. 

Fig. 61, ventral view of holotype; Fig. 62, lateral view; Fig. 63, holotype embedded in roof of mouth of fish; 
Fig. 64, another view of the same showing egg-sacs. 

Figs. 65-68. Lernaea longa, sp. nov. 

Fig. 65, lateral view of holotype showing anchor and swelling by legs 2. hd, head; Fig. 66. A paratype with the left 

ventral arm of the anchor distorted; Figs. 67 and 68, the externally visible parts of two other specimens embedded 

in the fish. The line near each figure is 1 mm. drawn to the same scale. 

The anchoring arms (Figs. 61 and 62) are four in number, of medium length and 
uniform thickness, each with a bend or a kink near the end. The ventral pair is 
directed slightly forwards and the dorsal pair backwards to the same extent. 

The head is not in line with the body, but inclined towards the angle between the 
ventral arms of the anchor. 

The egg-sacs are comparatively short and broad, being about 1-5 mm. long and 
0-5 mm. broad (Figs. 63 and 64). 



zoo. i, 1. 



18 ON SOME SPECIES OF LERNAE 

Lernaea longa sp. no v. 
Figs. 65-68 

Holotype, Reg. No. 1949.8. 14.27, and half a dozen paratypes, all females, in the 
British Museum. 

All the specimens were from a single specimen of Lates niloticus subsp. longispinus 
Worthington from Lake Rudolf, collected by Dr. E. B. Worthington in 1931. The 
parasites were embedded in the head and flanks of the fish. 

The length of the holotype is 19 mm., with the five pairs of limbs in positions 6-3, 
14, 36, 64, and yy per cent, of the body length from the anterior end. Owing to the 
fact that the head is held ventrally between the ventral arms of the anchor, these 
measurements have been made from the most anterior part of the body, i.e. the central 
boss of the anchor. In paratype, Reg. No. 1949.8. 14.29 (Fig. 66) the total length is 
22 mm. with the legs in positions 5, n, 32, 59, and 71 per cent, of the body length. 

The body is long and slender with a conspicuous swelling in the region of legs 2, 
and from this swelling a pair of rounded processes project ventrally with the second 
pair of legs between them. There are slight swellings in the regions of legs 3 and 4. 

Two examples will suffice to show how the torsion varies and may change its 
direction in this species. In the holotype the total torsion is a sinistral one of no°, 
made up of a dextral torsion of io° between legs 2 and 3 and a sinistral torsion 
between legs 3 and 4. In paratype Reg. No. 1949.8. 14.29 the total torsion is a dextral 
one of 20 ; this is the resultant of a sinistral torsion of 45 ° between legs 1 and 2, and 
of 135 ° between legs 2 and 3, followed by a dextral torsion of 90 between legs 3 and 
4, and of 110 between legs 4 and 5. 

The abdomen is very long, about a quarter of the total body length ; it is in line 
with the rest of the body and tapers gradually to a rounded tip without any indica- 
tions of segmentation. The pregenital prominence is ill defined. 

The anchor has normally four simple more or less cylindrical arms as shown in 
Fig. 65. One of the ventral arms of specimen Reg. No. 1949.8. 14.29 is branched, but 
this is evidently an abnormality probably caused by its meeting an obstruction during 
its growth into the flesh of the fish (Fig. 66). The ventral arms are about 5 times as 
long as they are broad and are directed backwards at an angle of about 45 ° to the 
body. The dorsal arms are a little shorter and are directed more nearly at right angles 
to the body. 

The head is placed on the ventral side of the anchor in the fork between the ventral 
pair of arms. 

None of the specimens had complete egg-sacs ; the most complete was 3 mm. long, 
with a maximum width of o-6 mm. 

r 

Lernaea haplocephala (Cunnington) 

1914 Lemaeocera haplocephala Cunnington, Proc. Zool. Soc. Lond. 1914: 826, pi. 1, figs. 4-7, 
text-fig. 1 b. 

1917 Lernaea haplocephala: Wilson, Proc. U.S. Nat. Mus. 53: 38. 

1918 ,, „ : Wilson, Bull. U.S. Bur. Fish. 35: 193, 195, pi. 15, fig. 92. 

1920 ,, ,, : Wilson, Bull. Amer. Mus. Nat. Hist. 43 (1) : 5, pi. 3, figs. 20-22. 



ON SOME SPECIES OF LERNAEA 19 

1923 Lernaeocera bichiri Kurtz, 5. B. Akad. Wiss. Wien. 131, Abt. 1: 332, pi. 2, figs. 1-11. 
1927 Lernaea haplocephala : Brian, Faune Colon. Fr. 1: 581, figs. 26-34. 
1944 ,, ,, : Capart, Bull. Mus. Hist. nat. Belg. 20 (24): 7. 

The British Museum possesses the twenty-seven specimens listed by Cunnington, 
from three species of Polypterus from Lake Tanganyika and the White Nile. I select 
as holotype the single specimen, Reg. No. 1914.12.2.3, taken from Polypterus congicus 
Boulenger collected from Lake Tanganyika by Cunnington himself and on which 
his description is largely based. L. haplocephala is probably the best known of the 
African species of Lernaea and has been found on several species of Polypterus in the 
White Nile, Belgian Congo, and Cameroons. 

The species is easily recognized by the shape of the anchor and by the peculiar 
swelling in the region of legs 2 ; and it is unfortunate that Wilson (1920), in his eager- 
ness to find characters in the appendages, should have added as a distinguishing 
character 'the small spherical terminal joint of the maxillipedes, with its four curved 
claws'. I have examined the maxillipedes of the holotype and of some of the para- 
types and can find no distinguishing feature in them ; they have five claws like every 
other species I have looked at. Wilson may have had a specimen with only four claws 
Brian describes and figures only three, but they are not easy to see and are difficult 
to count. There is, however, a character in which the appendages of L. haplocephala 
differ from those of all other species of Lernaea that I have been able to examine: 
there are only four setae on the terminal joints of the exopods of the first four pairs 
of legs (Table 2) ; other species have five setae here. The setation of these legs has 
been correctly figured by Kurtz and by Brian. 

Table 2 
Arrangement of the setae and spines on the legs of Lernaea haplocephala 





Leg 1 


Leg 2 


Leg 3 


Leg 4 


Exopod {^f 
End °P° d { setae' 


1 . 1 .2 
1. 1. 4 
0.0.2 
1. 1. 4 


1. 1. 3 

1. 1. 4 
0.0.2 
1.2.4 


1. 1. 3 

1. 1. 4 
0.0.2 
1.2.4 


1. 1. 3 

1. 1. 4 
0.0.2 
1.2.3 



Lernaea lophiara sp. nov. 

Figs. 69-80 

Holotype, Reg. No. 1949.8. 14.34, and several paratypes, all females, in the British 
Museum. The holotype was from the dorsal fin of Lethrinops lethrinus (Gunther) from 
Lake Nyasa. The paratypes included very similar specimens from the dorsal fins of 
the following species of fish, all from Lake Nyasa : Haplochromis prostoma Trewavas, 
H. sp. cf. micrentodon Regan, Rhamphochromis lucius Ahl, Pseudotropheus tropheops 
Regan, Diplotaxodon argenteus Trewavas, and also buried in the edge of the operculum 
of Lethrinops praeorbitalis Regan. Other paratypes which differ from the holotype 
only in having very short arms to the anchor were found in the dorsal fins of Haplo- 
chromis breviceps Regan and Tilapia melanopleura Dumeril. Specimens which I have 
left in situ and not examined but are presumably the same species were found in 



ON SOME SPECIES OF LERNAEA 



the dorsal fins of Haplochromis argyrosoma Regan, H. incola Trewavas, H. johnstoni 
(Giinther), and H. nigritaeniatus Trewavas. 

The length of the holotype is 9-6 mm. with the legs in positions 6, 16, 42, 76, and 
93 per cent, of the body length from the anterior end. The body is curved between 
legs 2 and 3, so that the axis of the anterior end of the body is approximately at 




Figs. 69-80. Lernaea lophiara sp. nov. 

Fig. 69, lateral view of head and anchor of holotype from dorsal fin of Lethrinops; Fig. 70, anterior view of the 
same; Fig. 71, paratype Reg No. 1949.8. 14.35 from dorsal fin of Haplochromis; Fig. 72, lateral view of anchor; 
Fig- 73, ventra j view of the same; Fig. 74, lateral view of abdomen and pregenital prominences of the same para- 
type; Fig. 75, ventral view of the same; Fig. 76, anterior view of anchor of paratype Reg. No. 1949.8. 14.45 from 
operculum of Lethrinops; Fig. 77, general view of this paratype; Fig. 78, posterior end with egg-sacs of paratype 
Reg. No. 1949.8. 14.46 from operculum of Lethrinops; Fig. 79, ventral view of abdomen and pregenital prominences 
of paratype Reg. No. 1949.8. 14.45; Fig. 80, lateral view of the same. 

Figs. 81-83. Lernaea cf. lophiara 

Fig. 81, specimen from operculum of Rhamphochromis Reg. No. 1949.8. 14.47; Fig. 82, ventral view of abdomen 
and pregenital prominences of this specimen; Fig. 83, lateral view of the same. The line near each figure is 1 mm. 

drawn to the same scale. 

right angles to the posterior end. The total torsion of the holotype is a dextral one 
of about 145 ; this is made up of a dextral torsion between legs 2 and 3 of 85 and 
a further dextral torsion of 6o° between legs 3 and 4. 

The lengths of seven paratypes range from 67 mm. to 9-8 mm., the positions 
of the five pairs of legs ranging from 5-6-5, 17-18, 42-47, 73-79, and 93-94 per cent, 
of the body length from the anterior end respectively. Only in two specimens, one of 



ON SOME SPECIES OF LERNAEA 21 

them the holotype, is the torsion of the body more than 90 . There is nearly always 
some torsion, however, and also a curvature in the region of legs 2 and 3. 

The anchor has four simple arms. In most of the specimens (Figs. 69, 70, 76, yy), 
including the holotype from the dorsal fin of Lethrinops praeorbitalis and paratype 
Reg. No. 1949.8. 14.45 from the operculum of the same fish, the dorsal arms are a 
little longer than the ventral ones and tend to splay outwards towards the ends. In 
two other specimens (Figs. 71-73) from dorsal fins, one from Haplochromis breviceps, 
the other from Tilapia melanopleura, the arms of the anchor are very short. 

The pregenital prominence is double in all specimens except one which is the 
smallest examined and measures 67 mm. in length. 

The setation of the legs is the same as that of L. cyprinacea (Table 1, p. 4). 

The egg-sacs are spindle-shaped and two or three times as long as the abdomen. 

Lernaea sp. cf. lophiara 
Figs. 81-83 

A fish of the species Rhamphochromis lucius Ahl bore three parasites : one on the 
fin is a typical example of Lernaea lophiara, the other two, one on the flank and 
the other on the operculum, are considerably larger than normal for that species 
and have very much larger arms to the anchor. The shape of the abdomen is also 
rather different. It may be found that these two specimens belong to a new species, 
but the comparatively well-developed condition of the arms of the anchor might be 
due to there being more space for them to grow in the body of the fish than there is 
in the fin. Against this, however, is the fact that the specimens of L. lophiara from 
the operculum of Lethrinops praeorbitalis do not differ in the size of the anchor or 
in other respects from specimens from the fins. 

The lengths of the two specimens from the flank and operculum are 14-4 mm. and 
137 mm. respectively. The anchor of the shorter specimen is damaged, that of the 
larger is shown in Fig. 81. The last segment of the abdomen is very small in these 
two specimens, the abdomen as a result being much more conical in shape than is 
typical for L. lophiara (Figs. 82-83). The positions of the five pairs of legs of the 
larger specimen are 5, 17, 46, 73, and 94 per cent, of the body length from the 
anterior end respectively. The setation of the legs is the same as that of the other 
specimens. 

Lernaea bistricornis sp. nov. 
Figs. 84-88 

Holotype Reg. No. 1949.8. 14.49 m tne British Museum. This species has to be 
described from a single specimen found at the base of a pelvic fin of Cardio-pharynx 
schoutedeni Poll from Lake Tanganyika. 

The length of the holotype is 87 mm. The body is curved evenly into a semicircle, 
and there is a sinistral torsion of about 90 . The positions of the five pairs of legs are 
8, 21, 45, 76, and 94 per cent, of the body length from the anterior end respectively. 
The setation of legs 1 and 2 cannot be seen, but that of legs 3 and 4 is the same as 
that of L. cyprinacea (Table 1, p. 4). 



22 



ON SOME SPECIES OF LERNAEA 



The anchor (Figs. 84-86) has six short, blunt processes, three on each side. There 
is a dorsal pair and a ventral pair, both of which are directed outwards and back- 
wards. These are similar to those of some specimens of L. lophiara ; but in addition 




Figs. 84-88. Lernaea bistricornis sp. nov. 

Fig. 84, latera Iview of anchor; Fig. 85, ventral view; Fig. 86, anterior view of the same; Fig. 87, ventral view of 
abdomen and pregenital prominence; Fig. 88, lateral view of the same. 

Fig. 89. Lernaea barbicola Leigh-Sharpe. Abdomen, pregenital prominence, and egg-sac. 

Figs. 90-95. Lernaea tuber osa sp. nov. 

Fig. 90, ventral view of holotype; Fig. 91, abdomen and pregenital prominence; Fig. 92, lateral view of anchor; 
Fig. 93, the same seen as a transparent object; Fig. 94, anterior view of anchor; Fig. 95, the visible part of the 
parasite protruding from the hole in the side of the fish. The line near each figure is 1 mm. drawn to the same scale. 

there is a pair of small knobs on each side of the head which reach over as if to 
protect it. 

The pregenital prominence is well defined (Figs. 87-88), but appears to be simple. 
The abdomen is of normal length, rather tapering, and without signs of segmentation. 

The egg-sacs are spindle-shaped and about 1-3 mm. long. 



ON SOME SPECIES OF LERNAEA 23 

Lernaea barbicola Leigh-Sharpe 
Fig. 89 

1930 Lernaea (Lernaeocera) barbicola Leigh-Sharpe. Parasitology 22: 334, text-figs. 1-6. 

Mr. Leigh-Sharpe has kindly presented the holotype Reg. No. 1949.8. 14.50 of this 
species mounted on a microscope slide to the Museum. It was from the tail of a species 
of Barbus from the Transvaal. 

Unfortunately the arms of the anchor have been broken and it is no longer possible 
to see their arrangement. 

Owing to the fact that the specimen is flattened on a slide the precise shape of the 
abdomen and pregenital prominence must remain uncertain; but I have made a 
camera lucida drawing (Fig. 89) which I hope is a little more accurate than Leigh- 
Sharpe's Fig. 2. Leigh-Sharpe' s figures of the first and second pairs of legs are 
evidently not intended to show the precise setation of these limbs. All four pairs 
are visible in the preparation, and with the help of an immersion lens I have been 
able to make out the setation, which is precisely the same as that of L. cyprinacea 
(Table I, p. 4). 

Lernaea tuber osa sp. no v. 
Figs. 90-95 

Holotype, Reg. No. 1949.8. 14.51, and one paratype in the British Museum. Both 
specimens were from the body of the fish Engraulicypris sardella (Giinther) from Lake 
Nyasa. The holotype was from the flank of a specimen collected by Dr. Christy in 
1925, and the paratype was from the mid- ventral line of a fish in the Museum 
collection, Reg. No. 1908.10.27.24-33 collected by Captain Rhoades. 

The length of both specimens is 11-5 mm. The positions of the five pairs of legs 
are 7-8, 18, 42, 72, and 91 per cent, in the holotype and y-8, 18, 44, 76, and 93 per cent, 
of the body length from the anterior end in the paratype. 

The total torsion in both specimens is 100 ° in a dextral sense. In the holotype this 
is the result of dextral torsions between legs 1 and 2 of 45 , between legs 2 and 3 of 40 , 
and between legs 3 and 4 of 15 . In the paratype the torsion is at first sinistral 
through 45 between legs 1 and 2, followed by dextral torsions of 105 ° between legs 
2 and 3, 35 between legs 3 and 4, and about 5 behind legs 4. 

The neck of both the specimens, that is the part of the body from the anchor to 
nearly the third legs, is covered with little peg-iike processes which immediately 
distinguish this species from any other known at present, and which have suggested 
to me the trivial name tuber osa. 

The anchor has four arms arranged as shown in Figs. 90 and 92-94 ; each bears a 
number of small finger-like processes. In spite of the apparent irregularity there is a 
distinct bilateral symmetry in the arrangement of the processes, and the two speci- 
mens are very similar to one another. 

The pregenital prominence is distinct but single or only indistinctly bilobed. The 
abdomen is of normal length and tapering, with the segments not marked off from 
one another. 

The egg-sacs of the holotype (Fig. 95) were small and rather shrunk in appearance. 



24 ON SOME SPECIES OF LERNAEA 

KEY TO ADULT FEMALES 

In the following key to the species of Lernaea those which I have seen are printed 
in bold type and those I know only from descriptions or figures are in italics. 
The following species are omitted as I consider them to be synonyms : 

L. bichiri (Kurtz, 1922) = L. haplocephala (Cunnington, 1914) 

L. carassii Tidd, 1933 = L. cyprinacea Linnaeus, 1758 

L. elegans Leigh-Sharpe, 1925 = L. cyprinacea Linnaeus, 1758 

L. pectoralis (Kellicott, 1882) = L. catostomi (Kr0yer, 1864) 

L. temnocephala (Cunnington, 1914) = L. barnimiana (Hartmann, 1865) 

L. tortua (Kellicott, 1881) = L. catostomi (Kr0yer, 1864) 

L. werneri (Kurtz, 1922) = L. composita Wilson, 1924 

1. Neck with many peg-like protuberances. . . L. tuberosa sp. nov. 
Neck smooth ........... 2 

2. Anchor with four unbranched arms, confluent at their bases ... 3 
Anchor with some other arrangement of its arms . . . ... 12 

3. A localized swelling at least twice the width of the body present in the 

region of legs 2 .......... 4 

Body not conspicuously swollen in region of legs 2 . . . . .5 

4. Abdomen of normal length, less than three times its breadth . 

L. haplocephala (Cunnington, 19 14) 
Abdomen very long, about a quarter of the total body length L. longa sp. nov. 

5. Arms of anchor long and straight and in a plane roughly parallel to body axis 6 
Arms not answering to this description ...... 7 

6. Pregenital prominence bilobed ..... L. tilapiae sp. nov. 
Pregenital prominence with three lobes . L. pomatoides (Kr0yer, 1864) 

7. Anchor with dorsal arms curved and ventral arms straight L. bagri sp. nov. 
Anchor not answering to this description ...... 8 

8. Abdomen short, little, if any, longer than pregenital prominence . . 9 
Abdomen distinctly longer than pregenital prominence . . . .10 

9. Dorsal and ventral arms of anchor of about equal size .... 

L. cruciata (Lesueur, 1824) 
Ventral arms much smaller than dorsal arms . L. tenuis (Wilson, 1916) 



L. lophiara sp. nov. 

. n 

L. palati sp. nov. 

composita Wilson, 1924 



10. Pregenital prominence bilobed 
Pregenital prominence simple 

11. Arms not tapering, each with a kink near the end 
Arms thick at base and tapering rapidly 

12. Anchor of four simple flattened arms, an anterior pair in front of legs 1 and 

a posterior pair behind these legs . . L. variabilis (Wilson, 19 16) 

Anchor not answering to this description . . . . . 13 

13. Main bulk of anchor at right angles to the body like the cross-bar of a T, 

with the length of cross-bar at least a third of body length . . 14 

Anchor not answering to this description . . . . . 17 

14. Anchor with a median dorsal process bifid at the tip ... . 

L. dolabroides Wilson, 19 18 



ON SOME SPECIES OF LERNA EA 25 

Anchor with no median dorsal process . . . . . . 15 

15. Lateral arms of anchor unbranched . . L. parasiluri Yamaguti, 1939 
Lateral arms of anchor each with a dorsal branch near the end . .16 

16. Anchor with a small pair of ventral arms near middle line, body without a 

conspicuous constriction . . . . . L. piscinae sp. nov. 

Anchor without ventral arms. Body with conspicuous constriction between 
legs 3 and legs 4 . . L. diceracephala (Cunnington, 1914) 

17. Anchor with six short, rounded protuberances, three on each side, a dorsal 

pair, a ventral pair, and also an anterior pair at the sides of the head 

L. bistricornis sp. nov. 
Anchor not answering to this description . . . . . .18 

18. Anchor set at right angles to body by a ventral flexure by legs 2, arms lateral 

with bulbous branches. Posterior part of body much swollen . 

L. insolens Wilson, 19 19 
Not answering to this description . . . . . . .19 

19. Anchor with a median dorsal arm which may be branched, and lateral arms . 20 
Anchor with arms in pairs, no median arm . . . . . .22 

20. Dorsal arm twice bifid, posterior half of body behind legs 4 swollen and 

spindle-shaped . . . . . . L. lagenula (Heller, 1865) 

Dorsal arm unbranched, or branched only once . . . . .21 

21. Lateral arms simple .... L. barbicola Leigh-Sharpe, 1930 
Lateral arms branched at least once . . . L. catostomi (Kr0yer, 1864) 

22. Arms of both dorsal and ventral pairs bifid . . L. oryzophila Monod, 1932 
Either dorsal or ventral arms unbranched . . . . . .23 

23. Ventral arms simple, dorsal arms branched . . . . . .24 

Ventral arms branched, dorsal arms unbranched . . . . .28 

24. Ventral arms very short, hardly longer than breadth of head . 

L. barnimiana (Hartmann, 1865) 
Ventral arms distinctly longer than breadth of head . . . -25 

25. Ventral arms curved outwards, with a small swelling facing anteriorly about 

the middle of the curve ...... L. barilii sp. nov. 

Ventral arms more or less straight, without a swelling . . . .26 

26. Legs 2 as well as legs 1 between the bases of ventral arms. Abdomen in line 

with the body .... L. ranae Stunkard & Cable, 1931 

Legs 2 situated some distance behind the bases of ventral arms. Abdomen 
generally at an angle with body . . . . . . .27 

27. Arms not more than three times as long as they are broad. Dorsal arms 

nearly as short as ventral ones. Egg-sacs oval L. esoscina (Burmeister, 1835) 
Arms slender and cylindrical in form. Dorsal arms distinctly longer than 
ventral. Egg-sacs spindle-shaped . . L. cyprinacea Linnaeus, 1758 

28. Branches of ventral arms unequal, main branch directed outwards and 

smaller one directed ventrally from it. Pregenital prominence bilobed . 

L. phoxinacea (Kr0yer, 1864) 
Ventral arms bifid at tip with resultant prongs equal and parallel. Pregenital 
prominence hemispherical. . . L. senegali Zimmermann, 1923 



26 ON SOME SPECIES OF LERNAEA 

REFERENCES 

Baird, W. 1850. The Natural History of the British Entomostraca. Ray Society, London. viii + 

364 pp., 36 pis. 
Barbut, J. 1783. The Genera Vermium. . . . London, xx+101 pp., n pis. 
Blainville, H. M. D. de. 1822. Memoire sur les Lern6es (Lernaea, Linn.). Journ. Phys. 95: 

372-380. 
Brian, A. 1927. Crustacea II. Copepoda parasitica. Faune Colon. Franc. 1: 571-587, figs. 1-34. 
1940. Sopra una specie di Copepodo parassita raccolto dal Prof. Parenzan nel lago Ararobi 

nell' A.O.I. Lernaea temnocephala (Cunnington) . Boll. Idrobiol. Caccia Pesca, 1: 50-56, 

text-figs. A-F. 
Burmeister, H. 1835. Beschreibung einiger neuen oder weniger bekannten Schmarotzer- 

krebse. . . . Nova Acta Leop. Carol. 17: 269-336, pis. 23, 24, 24 a, 25. 
Capart, A. 1944. Copepodes parasites des Poissons d'eau douce du Congo Beige. Bull. Mus. 

Hist. nat. Belg. 20 (24) : 1-24, text-figs. 1-4. 
Cunnington, W. A. 1914. Zoological results of the third Tanganyika Expedition, conducted 

by Dr. W. A. Cunnington, 1 904-1 905. Report on the Parasitic Eucopepoda. Proc. Zool. 

Soc. Lond. 1914: 819-829, pi. 1, text-fig. 1. 
Gurney, R. 1933. British Fresh-Water Copepoda, 3. Ray Society, London, xxix + 384 pp., 

text-figs. 1 196-2061. 
Hartmann, R. 1 865-1 866. Naturgeschichtlich-medicinische Skizze der Nilldnder. Berlin. 

pp. vii, 419. 2 abt. (abt. 2, pp. 2og~4ig, 1866.) 

1870. Beitrage zur anatomischen Kenntniss der Schmarotzer-Krebse. Arch. Anat. Phys. 

Wiss. Med. 1870: 726-752, pis. 17-18. 

1 87 1. "Dber das von Poren durchsetzte aussere Chitinskelet des Caliopus, Cecrops, und 

gewisser Lernaeoceren. 5. B. Ges. naturf. Fr. Berl. 1870: 60-61. 

Heller, C. (1865). Crustacea. Reise der osterreichischen Fregatte Novara um die Erde in 

den Jahren 1857, 1858, i8$g. Zool. Theil. 2 (8) : 1-280, pis. 1-25. 
Hofer, B. 1904. Handbuch der Fischkrankheiten. Stuttgart, xv + 359 pp., 18 pis., 222 text- 
figs. 
Kellicott, D. S. 1 88 1. Lerneocera tortua n.s. Proc. Amer. Soc. Micr. 3rd. Ann. Meeting, Detroit 

1880: 41-43, pi., figs. 1-3. 
1882. On certain crustaceous parasites of fresh-water fishes. Proc. Amer. Soc. Micr. $th 

Ann. Meeting, Elmira 1882: 75-78. 
Kroyer, H. 1863-1864. Bidrag til Kundskab om Snyltekrebsene. Naturhist. Tidsskr. Kjoben- 

havn (3), 2: 75-426, pis. 1-18. 
Kurtz, H. 1923. Zwei neue Arten von Lernaeocera aus dem Nil. 5. B. Akad. Wiss. Wien. 

(Abt. 1) 131: 327-337. p!s. 1-2. 
Leigh-Sharpe, W. H. 1925. Lernaea {Lernaeocera) elegans n. sp. A parasitic Copepod of 

Anguilla japonica. Parasitology, 17: 245-251, text-figs. 1-5. 
1930. Lernaea (Lernaeocera) barbicola n.sp. A parasitic Copepod of Barbus sp. from the 

Transvaal. Parasitology , 22: 334-337, text-figs. 1-6. 
Leseuer, C. A. 1824. On three new species of parasitic vermes belonging to the Linnean genus 

Lernaea. J. Acad. Nat. Sci. Philad. 3: 286-293, pi. 11. 
Linnaeus, C. 1746. Fauna Svecica. Stockholmiae. xxvi + 411 pp., 2 pis. 

1758. Sy sterna Naturae. Ed. X. Tom. 1, Holmiae. 824 pp. 

Markewitsch, A. P. 1934. Die Schmarotzerkrebse der Fische der Ukraine. Ann. Mus. zool. 

polon. 10: 223-249, pis. 44-45. 
1937- Copepoda parasitica der Binnengewdsser der U.S.S.R. (Akad. Wiss. Ukr. S.S.R.) 

Kiew, 222 pp., 27 pis, 10 text-figs. [Ukranian, Germ. Summary.] 
Matsui, Y., and Kumada, A. 1928. ' Ikari-Mushi ' (Lernaea elegans Leigh-Sharpe), a new para- 
sitic Copepod of Japanese Eel. /. Fish. Inst. Tokyo, 23: 101-107, pis. 5-7. 
Monod, T. 1932. Contribution a l'etude de quelques Copepodes parasites de Poissons. Ann. 

Parasit. hum. comp. Paris, 10: 345-380, text-figs. 1-23. 



ON SOME SPECIES OF LERNAEA 27 

Nakai, N. 1927. On the development of a parasitic copepod, Lernaea elegans Leigh-Sharpe, 

infesting on Cyprinus carpio L. /. Fish. Inst. Tokyo, 23: 39-59, pis. 2-4, text-figs. 1-7. 
Neresheimer, E. 1909. Die parasitischen Copepoden. In Brauer, Die Susswasserfauna 

Deutschlands, 11: 70-84, text-figs. 311-345. 
Okada, Y. K. 1927. Cop6pode parasite des Amphibiens. Nouveau parasitisme de Lernaea 

cyprinacea L. Annot. zool. Jap. 11: 185-187, text-figs. 1-2. 
Pesta, O. 1934. Krebstiere oder Crustacea. I. Ruderfusser oder Copepoda. Dahl, Die Tierwelt 

Deutschl. 29: 1-68, text-figs. 1-42. 
Scott, T. and A. 1913. The British Parasitic Copepoda. Ray Society, London, ix + 256 pp., 

2 pis. (Atlas: xii pp., 72 pis.) 
Stunkard, H. W. and Cable, R. M. 193 i. Notes on a species of Lernaea parasitic in the larvae 

of Rana clamitans. J. Parasit. 18: 92-97, pi. 8. 
Tidd, W. M. 1933. A new species of Lernaea (Parasitic Copepoda) from the Goldfish. Ohio J. 

Sci. 33: 465-468, pi. 1. 
Wagler, E. 1937. Crustacea (Krebstiere). Die Tierw. Mitteleuropas, 2, 2a: 3-224, text-figs. 

1-624. 
Wilson, C. B. 1916. Copepod parasites of fresh-water fishes and their economic relations to 

mussel glochidia. Bull. U.S. Bur. Fish. 34 [for 1914] : 331-374, pis. 60-74. 

191 7. North American Parasitic Copepods belonging to the Lernaeidae with a revision of 

the entire Family. Proc. U.S. Nat. Mus. 53: 1-150, pis. 1-2 1. 

191 8. The economic relations, anatomy, and life history of the genus Lernaea. Bull. U.S. 

Bur. Fish. Washington, 35 [for 1915-1916]: 163-198, pis. 6-15. 

19 1 9- A new species of parasitic copepod, with notes on species already described. Proc. 

U.S. Nat. Mus. 55: 313-316, pi. 21. 

1920. Parasitic Copepods from the Congo Basin. Bull. Amer. Mus. Nat. Hist. 43, 1 : 1-8, 

pis. 1-3. 

1924. Parasitic copepods from the White Nile and the Red Sea. Res. Swed. Zool. Exped. 

Egypt & White Nile, igoo-igoi. Pt. 5(3): 1-17, pis. 1-3. 

Yamaguti, S. 1939. Parasitic Copepods from Fishes of Japan. Pt. 5, Caligoida, III. Vol. Jubil. 

Prof. S. Yoshida, Osaka, 2: 443-487, pis. 14-33. 
Zimmermann, F. 1923. Bearbeitung der parasitischen Copepoden von Fischen. Denkschr. 

A had. Wiss. Wien. Math. Nat. Klasse, 98: ioi-iii, pis. 1-2, text-figs. 1-2. 




PRESENTED 

JAN ... 



AXISHSAINn 
3HX OX 
H3XNIHJ 

Aaxva saixvHO 

AS 

aaoaxo 
ssand AxisnaAiNn 

3HX XV 

nivxihs xvaao 
ni aaxNiHd 



1 APR 1950 

ON A GIANT SQUID 

OMMASTREPHES CAROLI Furtado 

STRANDED AT LOOE 

CORNWALL 

W. J. REES 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ZOOLOGY Vol.i No. 2 

LONDON : 1950 



ON A GIANT SQUID 

OMMASTREPHES CAROLI Furtado 

STRANDED AT LOOE 

CORNWALL 



BY 



W. J. REES, D.Sc. 




Pp. 29-42; Pis. 1-2; 12 text-figures; 3 map in the text 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ZOOLOGY V0I.1 No. 2 

LONDON : 1950 



THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted in 1949, is to be 
issued in five series, corresponding to the Departments 
of the Museum. 

Parts will appear at irregular intervals as they be- 
come ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

This paper is Vol. 1, No. 2, of the Zoological series. 



PRINTED BY ORDER OF THE TRUSTEES OF 
THE BRITISH MUSEUM 

Issued March 1950 Price Four shillings 



ON A GIANT SQUID, OMMASTREPHES CAROLI Furtado 
STRANDED AT LOOE, CORNWALL 

By W. J. REES, D.SC. 

The object of this note is to place on record some details of a female specimen 
of Ommastrephes caroli Furtado 1 stranded in live condition at Looe, Cornwall, in 
November 1940. It was acquired by the Plymouth Laboratory and was photographed 
before preservation by Mr. D. P. Wilson, to whom I am indebted for the excellent 
photographs. Subsequently it was preserved in formalin at the Laboratory, where 
I was able to examine it by kind permission of Mr. F. S. Russell, F.R.S. 

The earliest certain record of a stranding of this species, near Scheveningen in 
Holland in 1661, is mentioned by Steenstrup (1887), and in the same year the species 
was described for the first time by Furtado from Portuguese specimens in the Lisbon 
Museum. It was subsequently reported from the Faroes by Lonnberg (1897), and 
since that date there has been a number of records — all strandings — from British 
waters and one from Heligoland; these are summarized by Clarke & Robson (1929) 
and, more recently, by Stephen (1944). Apart from these positive records, there are 
occasional reports of strandings unsupported by details, and probably also strandings 
on lonely coasts which are never reported, so that the number of actual strandings is 
possibly much more frequent than indicated in the literature. 

It is curious that this species is known only from strandings and that all the known 
specimens are females. 0. caroli most nearly resembles 0. bartrami (Lesueur, 1821), 
from which it can be readily distinguished by the remarkable membranes of the 
third arms — this feature being absent in 0. bartrami and 0. pteropus. Robson (1925) 
described the largest example of 0. caroli yet found from a stranding at Withernsea, 
Yorkshire, and although the Looe specimen is a little smaller it is larger than all the 
others that have been measured. 

The standard measurements of 0. caroli from Looe are given below: 



Measurements in mm. 
Overall length (apex to tip of right tentacle) 
Total length (i.e. including 3rd arm) 
Dorsal mantle length .... 
Ventral mantle length 
Maximum mantle width (excluding fins) 
Maximum mantle width (including fins) 
Width at mantle openings . 
Length of head ..... 
Interocular width .... 
Thickness of head .... 
Arm length: 



,860 
,225 
670 
650 
245 
57° 
205 
170 
170 
100 



1st 

2nd ...... 

3rd 

4th 

Tentacle length .... 
Tentacle, length of sucker-bearing surface 



Right 
360 
415 
415 
445 

1,100 

463 



Left 
355 
415 
400 

445 

1,300 

460 



1 I have followed Winckworth (1932) in referring this species to Ommastrephes although most authors 
have recorded the species under the name Sthenoteuthis caroli. 



32 ON A GIANT SQUID, OMMASTREPHES CAROLI Furtado 

The Looe specimen agrees well with Robson's Withernsea example as regards colour 
and most external features and I have omitted further reference to them. I have, 
however, thought it desirable to redescribe the tentacles, arms, and suckers in some 
detail. 

The first pair of arms are quadrate in section and carry 25-26 pairs of suckers in 
oblique pairs on ridges. The proximal six pairs are well spaced, then distally, the 
remaining pairs are set closer together and give the appearance of being alternate. 
Suckers in the first or proximal row have a diameter of less than 10 mm. Those of the 
second to the eleventh rows are 10 mm. or over in diameter, while those of rows 
12-26 gradually decrease in size down to 1 mm. in diameter. On the right arm the 
largest suckers (on the fifth row) have a diameter of 13 mm. The left arm is very 
similar, with suckers of 14 mm. diameter in the fourth row. 

Both second arms are strongly keeled along their whole length and there are twenty- 
seven rows of suckers beginning with medium-sized proximal ones of 9 mm. in 
diameter. Distally there is a gradual increase in sucker width to 20 mm. in the eighth 
row, followed by an abrupt reduction to 12-15 mm. in the ninth row. 

The third arms have about twenty-eight pairs of suckers with similar appearance 
to those of the second arms. The proximal suckers are only 8 mm. in width, with a 
gradual increase distally to 13 mm. in the ninth row, followed by a gradual decrease. 
There is a well-developed keel which is much enlarged not far from the tip of the 
tentacle to form a strong crest. This is 70 mm. deep opposite the twenty-third and 
twenty-fourth rows of suckers. The lateral membrane, too, is very well developed 
and has a distinctive and characteristic shape — at least in the female, for the male is 
unknown. It extends from the base of the arm to within 60 mm. of its tip. The 
membrane is greatly enlarged distally to form a large, thin flap of a curious shape 
(Pis. 1 & 2). In the left arm this has a width of 220 mm., while in the right arm 
it is rather torn and is estimated to have a width in excess of 160 mm. Robson (1925) 
has discussed the shape of this organ in relation to the differentiation of species, but 
it is apparent, even in this fine specimen, that little reliance can be placed on it for 
taxonomic purposes because of its fragile nature. 

The right and left ventral arms have thirty-six and thirty-four pairs of suckers 
respectively ; these are widely spaced on the flat, sucker-bearing face of the tentacle. 
On the right arm the proximal suckers have a diameter of 7 mm., and there is a 
gradual increase in size to 14 mm. in the seventh row. Large suckers of 12-14 mm. 
diameter are maintained to the tenth pair, after which there is a gradual decrease 
down to 1 mm. or less at the tip of the arm. The left arm is similar, with larger 
suckers of 15-16 mm. diameter in the sixth to ninth rows. 

The right and left tentacles respectively are 1-65 and 1-94 times the length of the 
dorsal mantle. The following description applies to the right tentacle. It can be 
conveniently differentiated into four regions to facilitate description: viz. the tip 
portion, the large sucker region, the locking-apparatus region, and the proximal portion 
devoid of suckers. 

The tip portion, 87 mm. long, carries oblique rows of four suckers each at the 
extreme tip ; these are small with a diameter of 1 mm. Proximally these become 
enlarged to 5-6 mm. diameter with only three in a row. 



STRANDED AT LOOE, CORNWALL 33 

In the large sucker region of the manus there are eleven rows of suckers with four 
to each oblique row. The two median ones in each row are much enlarged, reaching 
a maximum size of 17-21 mm. ; those of the first and second row adjoining the tip 
portion are slightly smaller with diameters of 10 and 13 mm. respectively. On each 
side, flanking the median suckers, are smaller, long-stalked suckers of about 8 mm. 
diameter. These are borne on the transverse ridges. 

The locking-apparatus region (carpus) has three tubercules alternating with three 
smooth-ringed suckers and is similar in arrangement to that figured by Goodrich 
(1892) for 0. pteropus. These smooth carpal suckers are small with a diameter of only 
3 mm. The ordinary suckers of this region, counting from the most distal tubercule, 
are twelve in number and diminish in size down to 5 mm. proximally. 

On the sucker-less part of the tentacle there are fourteen transverse ridges which 
become fainter and disappear towards the base. 

The tentacle is keeled along its dorsal surface and becomes slightly finned in the 
part corresponding to the distal half of the large sucker region and the proximal half 
of the tip portion. There are narrow, undulating fins along both sides of the sucker- 
bearing face. Proximally the fin on the dorsal edge is less prominent but persists 
as a thin ridge as far as the end of the transverse ridges. The ventral fin reaches only 
to the tenth transverse ridge (from the base). Sucker rings of this species have been 
figured by Furtado and by Lonnberg, but unfortunately those of Furtado are not 
very clear and Lonnberg has failed to indicate the precise position of the suckers on 
the arms and tentacles. As Robson (1925) has pointed out, the dentition of the rings 
varies according to their position, the proximal teeth of the arm suckers being lost 
towards the free end of the arm. The earlier figures are therefore of little use for 
comparison, so new ones have been drawn from known positions on the arms 
(Figs. 1-3). 

On the basal portion of the arms the suckers are toothed all round, but the proximal 
teeth are small and often rudimentary (Fig. 3). Distal sucker rings have lost their 
proximal teeth and are of the form illustrated in Figs. 1 and 2. Typically these suckers 
have seven, long, back war dly directed teeth. The points where the proximal teeth 
disappear on each arm are fully discussed by Robson (1925). 

The tentacular sucker rings are dentate all round and also show some variation 
according to their position (Figs. 4-6). The distal teeth are curved inwards, while 
the proximal teeth, although often reduced in size, are bent outwards in the same 
direction as the distal ones; thus the teeth of the whole ring are admirably arranged 
for clawing. Fig. 4 illustrates a ring with twenty-one teeth, whereas that portrayed 
in Fig. 5 has twenty- three teeth. The larger rings of the manus are typically ommas- 
trephid in character with four enlarged teeth (one in each quadrant). This, the 
largest sucker ring of the club, has twenty-seven teeth. 

The stranding of giant squids of the genera Architeuthis and Ommastrephes on 
British coasts has aroused much interest during the past twenty years; the signi- 
ficance of the strandings, especially the preponderance of records along the east coast 
of Britain, being the subject of speculation by Clarke & Robson (1929), Robson 
(1933) > and Stephen (1944). 

The known strandings of 0. caroli, 0. pteropus, and Architeuthis spp. are plotted on 





Id 



lb 





2a 



2b 





3a 



3b 



Figs. 1-3. Sucker rings from the arms in face and oblique views. 

1 a & b., ring, 3-6 mm. in diameter, from 2nd left arm, 21st row. 

2 a & &., ring, 8-5 mm. in diameter, from 3rd left arm, 12th row. 

3 a & &., ring, 13 mm. in diameter, from 4^ left arm, 8th row. 





4b 





5a 



5b 





6a 



6b 



Figs. 4-6. Tentacular sucker-rings in face and oblique views. 

4 a & b., medium-sized sucker-ring, 3-75 mm. in diameter, from the tip portion of the tentacle. 

5 a & b., ring, 6-3 mm. diameter, from the long-stalked suckers of the manus. 

6 a & b., typical ring, 18 mm. in diameter, from the middle of the manus. 



36 ON A GIANT SQUID, OMMASTREPHES CAROLI Furtado 

Maps I— III, and it is at once evident that most of the specimens have come ashore 
at three places, viz. the Scarborough area, the Dunbar-North Berwick area, and at 
Buckie. Another feature of the strandings is that all, with the exception of a single 
record of 0. pteropus at North Berwick in June 1921, have come ashore during the 
winter months from November to March. 

Clarke & Robson correlate the strandings on the Yorkshire coast with hydro- 
graphic conditions which favour stranding, especially if the animal is enfeebled by 
some cause. They quote Bowman's testimony that a high percentage of drift bottles 
released in the north are finally stranded on the mid-Yorkshire coast and between 
Berwick and St. Abb's Head. 

Architeuthis and Ommastrephes are clearly oceanic species which occasionally 
migrate into the North Sea, possibly during the summer months, and are later en- 
feebled by unfavourable conditions during the winter months. There is as yet no 
clue as to what these factors are, but it is probable that lack of suitable food, lower 
salinity (especially near the coast), and temperature fluctuations have an adverse 
effect. 

Various Ommastrephids are, as young animals, common in the surface waters of 
temperate and tropical seas, but so far the habits of the large adults are a matter 
for speculation. Perhaps the single record of Ommastrephes pteropus (trawled off 
St. Kilda, at a depth of 180-200 fathoms in September 1925) is an indication of its 
normal habitat on the edge of the continental slope. Robson (1933) in discussing 
the distribution of Architeuthis was also inclined to favour this view. 

If we may judge by the records plotted on Maps I— III, 0. caroli is the most frequent 
immigrant into the North Sea, while 0. pteropus is just as rare as Architeuthis in 
British waters. 

The British records of these giant squids are scattered in the literature, and are, 
for the sake of completeness, given below. 

BRITISH RECORDS OF OMMASTREPHES CAROLI 

1. 8 Jan. 191 1. Briar Dene, Northumberland; Meek & Goddard (1926). Length (including 

3rd arm) 3 ft. 11 in. (1,175 mm.). 

2. Feb. 1921. Isle of Skye; Stephen (1944). 

3. 3 Jan. 1925. Withernsea, S. Yorkshire; Robson (1925). 

4. 7 Jan. 1925. Cullercoats, Northumberland; Meek & Goddard (1926). Length (including 

3rd arm) 3 ft. 8 in. (1,118 mm.). 

5. 14 Jan. 1927. Buckie, Moray Firth; Stephen (1944). 

6. March 1927. N. Berwick; Stephen (1944). 

7. 18 March 1927. N. Bay, Scarborough, Yorkshire ; Clarke & Robson (1929). Length 5 ft. 7 in. 

8. 1 Feb. 1928. Scarborough; Clarke & Robson (1929). Length (including 3rd arm) 3 ft. 6 in. 

9. Jan. 1929. Buckie, Moray Firth; Stephen (1944). 
10-11. Dec. 1929. N. Berwick; Stephen (1944), 2 specimens. 

12. 9 Jan. 1930. Filey, Yorkshire; Clarke (1930) & Stevenson (1935). Length (including 3rd 

arm) 3 ft. 9 in. 

13. 10 Feb. 1930. Isle of Skye; Stephen (1944). 

14. Feb. 1930. Isle of Mull; Stephen (1944). 

15. March 1930. Dunbar; Stephen (1944). 

16-17. 6 J an - I 93 I - Dunbar; Stephen (1944), 2 specimens. 

18. 22 Dec. 1931. South Sands, Scarborough; Stevenson (1935). Overall length 5 ft. 10 in. 



MAP I 



OMMASTREPHES 
CAROLI 




George Philip a. Son. Ltd 



The London Geographical Institute 



STRANDINGS OF OMMASTREPHES CAROLI FURTADO ON BRITISH COASTS 



ZOOL. I. 2 



MAP II 



OMMASTREPHES 
PTEROPUS 



? 




* Slrandinos 
O Other Records 



Georgf Philip & Son Lto 



The London Geographical Institute 



STRANDINGS AND OTHER RECORDS OF OMMASTREPHES PTEROPUS STEENSTRUP IN 

BRITISH WATERS 



MAP III 



ARCHITEUTHIS 
spp. 




# Stran&mds 
O Other Records 



George Philip & Son Ltd The London Geographical Institute 

STRANDINGS AND OTHER RECORDS OF ARCHITEUTHIS SPP. IN BRITISH WATERS 



4 o ON A GIANT SQUID, OMMASTREPHES CAROLI Furtado 

19. 12 Dec. 1932. Buckie, Moray Firth; Stephen (1933). Overall length 6 ft. 2 in. 

20. 31 Jan. 1935. South Bay, Scarborough; Clarke & Stevenson (1935). Overall length 5 ft. 

21. 13 Feb. 1935. 1 \ miles north of Scarborough; Clarke & Stevenson (1935). Overall length 

5 ft. 2 in. 

22. 3 Nov. 1935. Castlerock, Co. Londonderry; Stendall (1936). Determined by A. C. Stephen. 

23. 24 Nov. 1937. Birsay Parish, Orkney; Stephen (1938). Overall length 5 ft. 

24. 18 Dec. 1937. Stronsay, N. Orkney; Stephen (1938). Overall length 5 ft. 8 in. 

25. Nov. 1940. Looe, Cornwall (present record) . 

26. Jan. 1941. Fair Isle, Shetland; Stephen (1944). 

BRITISH RECORDS OF OMMASTREPHES PTEROPUS STEENSTRUP 

1. 19 Nov. 1883. Scarborough; Goodrich (1892). 

2. 27 Feb. 1884. 'North Sea'; Goodrich (1892). 

3. Jan. 1892. Salcombe, Devon; Goodrich (1892). 

4. ? Killala, Co. Mayo; Nichols (1905, 'many years ago'). 

5. ? Miltown Malbay, Co. Clare; Nichols (1905, 'a few years ago'). 

6. 19 Dec. 1907. Redcar; Hoyle (1908). 

7. 1 Mar. 1912. Redcliff, near Scarborough. Length (including 3rd arm) 3 ft. 

8. June 1921. N. Berwick, Firth of Forth; Ritchie (1922). 

9. ? Isle of Man ; Robson & Chadwick MS. 

10. Sept. 1925. Trawled off St. Kilda in 180-200 fathoms. Overall length 6 ft. (det. Robson). 

BRITISH RECORDS OF ARCHITEUTHIS SPP. 

1. 1673. Dingle Bay, Co. Kerry, S. Ireland; (More, 1875: 4526, as Dinoteuthis proboscideus) . 

2. 1860-1861. Between Hillswick and Scalloway, W. Shetland ; (Jeffreys, 1869: 124, as Archi- 

teuthis monachus). 

3. 25 Apr. 1875. Caught at sea off Boffin Island, Connemara, Ireland; (More, 1875: 123). 

4. Oct. 1880. Stranded at Kilkee, Co. Clare, S. Ireland; (Ritchie, 1918: 137, as Architeuthis). 

5. 1 914. In stomach of a sperm whale at Belmullet Whaling Station; (Hamilton, 1915: 137). 

6. 2 Nov. 1917. Stranded at Dunbar, Firth of Forth; (Ritchie, 1918: 133, as Architeuthis 

harveyi) . 

7. Feb. 1920. Stranded at N. Uist, Outer Hebrides; (Ritchie, 1920: 57, as Architeuthis 

harveyi) . 

8. 192 1. Stranded at Caithness, Scotland; (Ritchie, 1922: 423, as Architeuthis harveyi). 

9. 14 Jan. 1933. Stranded at Scarborough, Yorkshire; (Robson, 1933, as Architeuthis clarkei 

n. sp.). 
10. 7 Nov. 1937. Off Bell Rock, Angus, E. Scotland; (Stephen, 1937, as Architeuthis harveyi). 

REFERENCES 

Clarke, W. J., & Robson, G. C. 1929. Notes on the stranding of giant squids on the north-east 

coast of England. Proc. Malacol. Soc. Lond. 18 : 154-158 ; 1 text-fig. 

& Stevenson, J. A. 1935. Yorkshire Cephalopods. /. Conch. 20 : 102. 

Daniel, R. J. 1925. A large oigopsid cephalopod. 39th Ann. Rep. Mar. Biol. Stat. Port Erin, 

1925 : 34 i 1 text-fig. 
Furtado, A. 1 887. Sur une nouvelle espece de C6phalopode appartenant au genre Ommatostrephes. 

Mem. R. Acad. Lisboa, 6 (2): 1-16; 2 pis., 5 text-figs. 
Goodrich, E. S. 1892. Note on a large squid (Ommastrephes pteropus Steenstrup). /. Mar. Biol. 

Assoc. U.K., n.s., 2: 314-321; text-figs. 
Grieg, J. A. 1933. Cephalopods from the west coast of Norway. Bergens Mus. Aarb. 1933 (4) : 

1-25 ; pis. 1-4, 1 text-fig. 
Grimpe, G. 1925. Zur Kenntnis der Cephalopoden-Fauna der Nordsee. Wiss. Meeresuntersuch. 

16 (3) : 1-124; 1 pi., 34 text-figs. 



STRANDED AT LOOE, CORNWALL 41 

Hamilton, J. E. 1915. Belmullet Whaling Station: Report to the Committee. Rep. Brit. Ass. 

1914: 1 25-1 61 ; 4 text-figs. 
Hertling, H. 1938. Ueber eine auf Juist gestrandete Sthenoteuthis caroli (Furtado). Wiss. 

Meeresuntersuch. 1 : 93-1 11. 
Hoyle, W. E. 1908. A large squid at Redcar. Naturalist, 615: 132-133; 1 text-fig. 
Jeffreys, J. G. 1869. British Conchology, 5. 
Lonnberg, E. 1897. Ofversigt ofver Sveriges Cephalopoder. Bih. Svensk. Vetenskakad. Handl. 

17 (4, No. 6) : 1-41 ; 1 pi. 
Massy, A. L. 1928. The Cephalopoda of the Irish coast. Proc. R. Irish Acad. 38 (B, No. 2) : 25-37. 
Meek, A., & Goddard, T. R. 1926. On two specimens of giant squid stranded on the Northum- 
brian coast. Trans. Nat. Hist. Soc, Northumb., n.s., 6 : 229-237. 
More, A. G. 1875 a. Gigantic squid on the west coast of Ireland. Ann. Mag. Nat. Hist. (4), 

16: 123-124. 
1875 b. Notice of a gigantic cephalopod (Dinoteuthis proboscideus) , which was stranded at 

Dingle, in Kerry, two hundred years ago. Zoologist, 83 : 4526-4532. 
Nichols, A. R. 1905. On some Irish specimens of a large squid, Sthenoteuthis pteropus (Steen- 

strup). Irish Nat. 14: 54-57; 1 text-fig. 
Ritchie, J. 191 8. Occurrence of a giant squid (Architeuthis) on the Scottish Coast. Scot. Nat. 

1918: 133-139. 

1920. Giant squid cast ashore N. Uist, Outer Hebrides. Scot. Nat. 1920 : 57. 

1922. Giant squid on the Scottish coast. Rep. Brit. Ass. 1921 : 423. 

Robson, G. C. 1925. On a specimen of the rare squid, Sthenoteuthis caroli, stranded on the 

Yorkshire coast. Proc. Zool. Soc. Lond. 1925 : 291-301 ; pi. 1 ; 5 text-figs. 
1933- Ori Architeuthis clarkei, a new species of giant squid, with observations on the genus. 

Ibid. 1933 : 681-697 ; pi. 1, 8 text-figs. 
Stendall, J. A. S. 1936. Giant cuttlefish, Sthenoteuthis caroli Furtado, ashore in Co. London- 
derry. Irish Nat. J. 6 : 23-24. 
Stephen, A. C. 1933. Rare cuttlefish {Sthenoteuthis caroli) washed ashore at Buckie. Scot. Nat. 

1933 : 96. 

1938. Rare squid in Orkney. Ibid. 1938: 119. 

1944. The Cephalopoda of Scottish and adjacent waters. Trans. Roy. Soc. Edinb. 61 : 

247-270; 14 text-figs. 
Stevenson, J. A. 1935. The Cephalopods of the Yorkshire coast. /. Conch. 20: 104-116; 

pis. 3-7, 1 text-fig. 
Verrill, A. E. 1879-1881. The Cephalopods of the north-eastern coast of America. Trans. Conn. 

Acad. Arts. Sci. 5: 177-257 and 259-446; pis. 13-56. 
Winckworth, R. 1932. The British Marine Mollusca. /. Conch. 19: 211-252. 



PRESENTED 

1 ai ;. . 



B.M. (N.H.) Zoology, I, 2 



PLATE 1 




Photo. D. P. Wilson 



OMMASTREPHES CAROLI ; DORSAL VIEW 



PLATE 2 




Photo. D. P. Wilson 



OMMASTREPHES CAROLI ; VENTRAL VIEW 




PRESENTED 

1 APR 1950 






PRINTED IN 

GREAT BRITAIN 

AT THE 

UNIVERSITY PRESS 

OXFORD 

BY 

CHARLES BATEY 

PRINTER 

TO THE 
UNIVERSITY 




- 1 APk i960 

THE^*0ENTITY OF 

CAPTAIN COOK'S 

KANGAROO 



T. C. S. MORRISON-SCOTT 

AND 

F. C. SAWYER 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ZOOLOGY Vol. i No. 3 

LONDON : 1950 



THE IDENTITY OF 
CAPTAIN COOK'S KANGAROO 



BY 

T. C. S. MORRISON -SCOTT 

AND 






F. C. SAWYER 



"H 



Pp. 43-50; Ph. 3-5 




BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ZOOLOGY Vol. 1 No. 3 

LONDON: 1950 



THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted in ig4g, is to be 
issued in five series, corresponding to the Departments 
of the Museum. 

Parts will appear at irregular intervals as they be- 
come ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

This paper is Vol. i, No. 3, of the Zoological series. 



PRINTED BY ORDER OF THE TRUSTEES OF 
THE BRITISH MUSEUM 

Issued March ig$o Price Three shillings 



THE IDENTITY OF CAPTAIN COOK'S KANGAROO 

By t. c. s. morrison-scott and F. c. sawyer 

INTRODUCTION 
The identity of the kangaroo discovered by Captain Cook's expedition in 1770 has 
lately been the subject of some dispute. For years this kangaroo has been referred 
to as Macropus giganteus (Zimmermann, 1777) and was thought to have been the 
Great Grey Kangaroo, until Iredale & Troughton (1925) not only pointed out that 
giganteus is antedated by Mus canguru Miiller, 1776, which was based on the descrip- 
tion and plate given in Hawkesworth's (1773) account of Cook's voyage, but also 
threw doubt on whether Captain Cook's kangaroo was in fact the Great Grey 
Kangaroo. 

The ship's company of H.M.S. Endeavour included Sir Joseph Banks who brought 
with him Dr. Solander as naturalist and Sydney Parkinson as draughtsman. Iredale 
& Troughton (1925) published a transcript of Solander's manuscript Latin description 
of the kangaroos obtained by Captain Cook's party at Endeavour River (the future 
site of Cooktown) in June and July 1770 — a description which, as Iredale & Troughton 
pointed out, does not accord too well with the Great Grey Kangaroo. They sup- 
ported their contention that the animals in question were not Great Grey Kangaroos 
with the statement that the latter do not occur at or near Cooktown. Even if this 
were true the argument would not be valid, since the non-occurrence of the species 
at Cooktown nowadays does not preclude its possible occurrence there in 1770, when 
the country was quite undeveloped. But in fact Raven (1939) records that the Great 
Grey Kangaroo occurred within thirty miles of Cooktown in 1897, and Tate informs 
us (in litt.) that he obtained three specimens about fifteen miles from Cooktown in 
1947. 

Iredale & Troughton, though satisfied in their own minds that Captain Cook's 
kangaroo is not conspecific with the Great Grey Kangaroo, were unable to decide its 
identity but suggested that the weight of evidence pointed to a form of the robustus 
series. 

The same authors (1937) next published a paper in which they sought to show that 
Captain Cook's kangaroo was a northern representative of the Whiptail, or Pretty- 
face Wallaby — usually known as Macropus (Protemnodon) parry i Bennett, but which 
they hold should be called Wallabia elegans Lambert. This contention rests on rather 
insecure foundations. Briefly, the argument is that in 1929, or thereabouts, two skins 
I were purchased in the neighbourhood of Cooktown and that Solander's description, 
they say, agrees with one of these which was a Whiptail — the other skin being that of 
a Wallaroo of the antilopinus type. But it is not at all clear why Cook's kangaroo 
must necessarily be restricted to one of the two species represented by these two 
! purchased skins, nor is it clear why Iredale & Troughton abandoned their previous 
iconclusions that the weight of evidence pointed to Cook's kangaroo having been 
a form of robustus. 



46 THE IDENTITY OF CAPTAIN COOK'S KANGAROO 

The next stage in the controversy was a paper by Raven (1939), who holds that the 
evidence is decidedly against Cook's kangaroo having been a Whiptail, or Pretty-face 
Wallaby. With this the present writers concur. Raven further holds that the evi- 
dence supports the view that the early revisers were right in identifying Cook's 
kangaroo with the Great Grey Kangaroo and pleads the confusion caused by up- 
setting this position. 

Finally Tate (1948), in the course of his review of the Macropodidae, dismissed the 
Whiptail theory of Iredale & Troughton and agreed with Raven that, inter alia, the 
hip stripe and face stripe of the Whiptail are too diagnostic to have been omitted 
from the contemporary plate and descriptions of Cook's kangaroo had the animal in 
fact been a Whiptail. Tate added that the only really large species of Macropodidae 
that conceivably could have been found near Cooktown are the Great Grey, the Red, 
and Macropus robustus reginae Schwarz, one of the antilopine group. He dismissed 
the second and third on grounds of colour and decided that the description and plate 
in Hawkesworth (1773) — and hence Captain Cook's kangaroo and Mus canguru 
Miiller, 1776 — agreed most closely with the Great Grey Kangaroo. 

Tate avoided discussion of Solander's manuscript description, but since Solander 
was on board Cook's ship in his capacity as a naturalist, what he has said on the 
subject of the kangaroos must be examined. Here, however, we are straightway 
confronted with a difficulty. 

Iredale & Troughton (1937) say that Solander's description was based upon the 
small male first captured, and Troughton (1946: 202) repeats the contention, saying 
that it is indisputable that it applies only to an apparently adult male weighing 
38 pounds. But far from being indisputable it is not at all clear why these authors 
take this view at all, unless it is because the only measurements given are those of the 
male which Mr. Gore shot on 14 July 1770 (Solander gives the weight of this animal 
as 24 pounds ; the difference between this and the 38 pounds of the other accounts 
may be the difference between the 'clean' and 'dead weight'). But Solander gives 
the weights of all three animals taken, and the description itself is clearly a composite 
one since both male and female genitalia are described and also the mammae, and 
Solander says that the size of the animal varies with age. Nor is it clear why Trough- 
ton refers to the 38-pound animal as apparently adult when Solander says that it was 
possibly two or three years old. Solander's estimate of its years may not be reliable, 
but he was basing his view that it was not adult on the condition of the molar teeth, 
as will be seen from his discussion of the latter. 

But on top of this, Solander may well have had three separate species as well as 
three separate specimens in front of him as he wrote, and it is not possible to say 
which animal he had most in mind while describing the various characters. He might 
well have been making a qualitative average of the characters of all three. So Solan- 
der is not much help in arriving at the identity of Cook's kangaroo and any deductions 
drawn from his description should be treated with reserve. With this in mind it can 
be said that in two particulars Solander's description does not encourage any leanings 
towards the Great Grey theory. The rhinarium is described as 'Rostrum breviusculum, 
parum compressum ; apice inter nares nudum ibique cute aterrima rugulosa vestitum'. 
But the Great Grey Kangaroo has hairy skin between the nostrils. Then again the 



THE IDENTITY OF CAPTAIN COOK'S KANGAROO 47 

upper incisors are described as ' Incisores sex, approximati, lati : primum par leviter 
bilobum; secundum integrum; tertium latius crassiusque, bilobum: lobis anticis 
minoribus '. Iredale & Troughton on the one hand, and Raven on the other, perform 
some agile juggling with the Latin text in support of their respective theses, but what 
Solander says is that the third upper pair of incisors are bilobed and that the anterior 
lobes are the smaller, thus suiting neither the Whiptail theory nor the Great Grey 
theory. However, as has already been indicated, Solander's description cannot be 
treated as a reliable guide in the quest for Captain Cook's kangaroo. 

The controversy has so far been argued in terms of Solander, Hawkesworth, and a 
skin obtained near Cooktown 160 years after Cook was there. It seems strange that 
no attempt appears to have been made to find the original specimens, especially as 
Iredale & Troughton (1925), quoting Hunter (1790), drew attention to the probability 
of a skull which Banks gave to Hunter being in the Museum of the Royal College of 
Surgeons. Iredale & Troughton also drew attention to the probability of a pencil 
drawing by Parkinson being preserved in the British Museum. But they did not 
pursue these two lines of research on which we now report. 

DRAWINGS BY SYDNEY PARKINSON AND NATHANIEL DANCE 
It seems certain that the plate of Captain Cook's kangaroo published by Hawkes- 
worth (1773) was based on a drawing by Sydney Parkinson, the draughtsman in 
Banks's employ on board H.M.S. Endeavour. Search has been made in the British 
Museum (Natural History), and though the original of Hawkesworth's plate has not 
been found there are two rough sketches of kangaroos signed 'S. Parkinson' and 
marked in his hand 'Kangura Endeavour's River'. On the back of one of these 
Parkinson has added, The whole body pale ash colour the ears excepting the base 
fine specled gray iris of the eye Chestnut '. It was the practice of Parkinson, and other 
artists who accompanied Cook on his voyages, to make pencil sketches of animals 
seen, together with notes on the details of coloration, &c, the intention being to 
paint these in at a later date. In Parkinson's case, due to his death before the end of 
the voyage, many of the sketches were never completed. These drawings are inade- 
quate for purposes of identification but we consider them of sufficient interest to 
warrant publication (PI. 3). 

Of much greater interest, however, is a wash drawing of a complete kangaroo skull 
and another of its lower jaw shown separately (PL 4). These are signed 'N. Dance' 
and are among the collection of Parkinson drawings which came to the British 
Museum from Sir Joseph Banks's library. Dryander (1748-1810), in his manuscript 
catalogue of the drawings of animals in Banks's library, has the following entry on 
page 21 : 

Mammalia — Glires, Kanguru 

— K N.C. S. Parkinson 

x Cranium Nath. Dance 

The ' — 'is Dryander's symbol for a pencil drawing and the 'x' for a coloured one; 
' N.C. ' stands for Nova Cambria, as that part of Australia was called in those days. 
Sir Nathaniel Dance (1735-1811) was a celebrated portrait painter with a reputa- 



48 THE IDENTITY OF CAPTAIN COOK'S KANGAROO 

tion for accuracy as a draughtsman. Captain Cook sat to him for his portrait in 1776 
(fide Kitson, 1907), after which year Dance appears to have given up painting. 

There is no indication of the scale of the drawing, but by analogy with the series of 
Parkinson drawings it seems likely that the skull is drawn life-size. The skull and 
lower jaw are both represented on a single folio sheet. Parkinson's drawings are also 
on folio sheets and his practice was to draw objects life-size except where they were 
too big for the paper. In this case he reduced them, but he did not make drawings 
larger than life-size. This seems to have been the general practice of the time. The 
point is not pressed, but if Dance's drawing is life-size, then it is likely to be that of 
the skull of the 84-pound kangaroo shot 1 on 27 July 1770 ; the other two beasts, one 
shot by Lieutenant Gore on 14 July and another caught by Banks's greyhound on 
29 July, were smaller. 

The skull drawn by Dance appears to be that of a young Macropus robustus. We 
have been unable to trace the skull itself. 

ANOTHER OF CAPTAIN COOK'S SPECIMENS 
John Hunter, in his observations on animals in White's Journal (1790), says: 'Of 
the Kangaroo . . . the only parts at first brought home were some skins and sculls ; 
and I was favoured with one of the sculls from Sir Joseph Banks.' The posthumous 
papers of Hunter (1728-93) edited by Owen (1861) contain the same words, but Owen 
has added a footnote to the last sentence quoted above, which reads: 'No. 1732 
Hunt. Osteol.' 

Professor Wood-Jones has searched for this skull in the Museum of the Royal 
College of Surgeons but it cannot be found, and appears to have been destroyed by 
bombs along with many other Hunterian specimens during the 1939-45 War. How- 
ever, he drew our attention to a figure of a skull in a paper on the history of surgery 
by Webb- Johnson (1939). The text to this figure says: 'Kangaroo's skull, from the 
Hunterian collection brought from Australia ("New Holland") by Sir Joseph Banks 
when with Captain Cook's Expedition, 1768-71.' The figure itself is a reproduction 
of a photograph and it shows quite clearly the number ' 3703 ' painted on the skull. 
Flower's catalogue (1884) makes it plain that No. 3703 is the same specimen as No. j 
1732 in Owen's catalogue (1853). Webb- Johnson's figure is small and not very 
clear, but Professor Wood- Jones went to much trouble and eventually found a 
lantern slide of the same photograph. This slide (PL 5) is probably one Webb- 
Johnson had made when he read his paper in 1939. The skull it represents is clearly 
not the same as the one drawn by Dance and it appears to be slightly younger, an 
impression which is borne out by the description of its dentition in Owen's catalogue 
(1853). The skull is from one of the three animals obtained at Endeavour River in 
1770 and is probably that of the 38-pound animal shot by Lieutenant Gore on 
14 July 1770. 

As will be seen from Plate 5, the skull is a young one and the incisors are missing. 
In view of its important bearing on the nomenclature of the genus Macropus, we sent 
the photograph to Dr. G. H. H. Tate, who has recently (1948) monographed the 

1 By Lieutenant Gore, according to the journal of Midshipman John Bootie, who records the weight 
as 80 pounds. 



THE IDENTITY OF CAPTAIN COOK'S KANGAROO 49 

kangaroos, and himself collected specimens in the neighbourhood of Cooktown. We 
are indebted to him for his detailed report on this skull which he unhesitatingly refers 
to the Great Grey Kangaroo — amongst other characters the short ante-orbital canal 
and the ' zog ' in the maxillo-premaxillary suture being particularly characteristic. 

CONCLUSION 

Captain Cook's first expedition to Australia obtained three specimens of kangaroo, 
all from Endeavour River, Queensland, July 1770. The skull of one of these was still 
preserved in the Museum of the Royal College of Surgeons in 1939 but was destroyed 
by bombs during the late war. No trace of the other material has been found. 

The only figure of the original material hitherto generally known to zoologists is the 
plate in Hawkesworth (1773) of a not easily determinable kangaroo, or reproductions 
of it. Four more figures are now published. The first two are indeterminable outline 
drawings of the whole animal by Parkinson, who was on board Cook's ship. The third 
is a painting of a skull by Nathaniel Dance. This is almost certainly the skull of one 
of Cook's specimens ; in fact it is difficult to see where else it could have come from. 
It is the skull of a Wallaroo of the Macropus robustus series. 

The fourth is a photograph (PL 5) of the specimen which was destroyed in the 
Museum of the Royal College of Surgeons. This skull was from one of the kangaroos 
obtained by Cook's party at Endeavour River in July 1770. It was given by Banks 
to Hunter and is No. 1732 in Owen's catalogue (1853) and No. 3703 in Flower's 
catalogue (1884). It is the skull of a young Great Grey Kangaroo and we hereby 
designate it as the photo-lectotype of Macropus canguru (Muller, 1776) — ' Captain 
Cook's Kangaroo '. 

REFERENCES 

Banks, J. 1768-1771. Journal. [MS. transcript, preserved in the Botanical Department, British 

Museum, by the Misses Mary and Hannah Turner, aunts of Sir J. D. Hooker who helped to 

collate it with the original which is now in the Mitchell Library, Sydney.] This MS. contains 

material which was omitted by Hooker in his published version of the Journal (1896). 
Barton, G. B. 1893. Historical Records of New South Wales, 1 (1) : 1-526. [This contains reprints 

of Cook's log, of the journals of his officers, including that of Midshipman J. Bootie, and of 

correspondence between Cook and the Admiralty and others concerning his voyages.] 
Bootie, J. 1770. See Barton, G. B. 
Carrington, H. 1939. Life of Captain Cook. London (Sidgwick & Jackson). [This contains a 

list of all the known MS. logs and journals of Cook, Banks, and other members of the ship's 

company of H.M.S. Endeavour.'] 
Cook, J. 1 768-1 771. Captain Cook's Journal during his First Voyage. . . . [Edited by Capt. 

W. J. L. Wharton, London, 1893, from a contemporary MS. transcript of Cook's holograph. 

This transcript is now in the Australian Museum, Sydney. Cook's holograph is in the 

National Library, Canberra.] 
Dryander, J. {Autograph Catalogue of the Drawings of Animals in the Library of Sir J. Banks, 

arranged in systematic order.] (Preserved in the Zoological Department, British Museum.) 
Flower, W. H. 1884. Catalogue of Specimens . . . in the Museum of the Royal College of Surgeons 

of England, 2 : 708. 
Hawkesworth, J. 1773. An Account of the Voyages undertaken . . .for making Discoveries in the 

Southern Hemisphere . . .by . . . Captain Cook, 3 : 577 (1st edition) ; 173 (2nd edition) . London. 
Hunter, J. 1790. See White, J. 



50 THE IDENTITY OF CAPTAIN COOK'S KANGAROO 

Hunter, J. 1861. Essays and Observations on Natural History, Anatomy, Physiology, Psychology 

and Geology, 2 1 250. [Edited posthumously by R. Owen.] 
Iredale, T., & Troughton, E. Le G. 1925. Captain Cook's Kangaroo. Aust. Zool. 3 : 311. 

1937. The identity of Cook's Kangaroo. Rec. Aust. Mus. 20 : 67. 

Kitson, A. 1907. Captain James Cook, &c. London (John Murray). Pp. xvi, 525, frontis., 

16 pis., 1 map. 
Muller, P. L. S. 1776. Des Ritters C. von Linne . . . vollstdndiges Natursystem nach der zwolften 

Lateinischen Ausgabe . . . Supplementsband : 62. Niirnberg. 
Owen, R. (1853). Descriptive Catalogue of the Osteological Series contained in the Museum of the 

Royal College of Surgeons of England, 1 : 322. London. 

1861. See Hunter, J. 

Parkinson, Stansfield. 1773. A Journal of a Voyage to the South Seas in His Majesty's Ship 

the Endeavour. Faithfully transcribed from the Papers of the late Sydney Parkinson, &c. : 145. 

London. [Sydney Parkinson died on 26 January 1771.] 
Parkinson, Sydney. 1 768-1 771. [294 Original Water Colour Drawings and Pencil Sketches of 

Animals made during Cook's First Voyage by S. Parkinson, and {ij of Fish and 4 of Mollusca) 

by A . Buchan.] 3 vols. 
Raven, H. C. 1939. The identity of Captain Cook's Kangaroo. /. Mammal. 20 : 50. 
Solander, D. C. 1 768-1 771. [Manuscript descriptions of Animals written on slips and systematically 

arranged in accordance with Linne' s ' Sy sterna Naturae . . . Editio duodecima reformata'.] 

1 : 90. (Preserved in the Zoological Department, British Museum.) 
Tate, G. H. H. 1948. Studies on the anatomy and phylogeny of the Macropodidae (Marsupialia) . 

Bull. Amer. Mus. Nat. Hist. 91: 233. 
Troughton, E. Le G. 1942. The kangaroo family — origin and earliest discoveries. Aust. Mus. 

Mag. 8: 17. 
1946. Furred Animals of Australia. (Third revised edition.) Pp. xxx, 376, 25 pis. Sydney 

(Angus & Robertson). 
Webb-Johnson, A. 1939. The George Adlington Syme Oration: Surgery in England in the 

making. Aust. N.Z. J. Surg. 9: 10. 
White, J. 1790. Journal of a Voyage to New South Wales &c. [Hunter discusses the kangaroo 

on p. 272.] London. 
Zimmermann, E. A. W. 1 777. Specimen zoologiae geographicae, Quadrupedum domicilia et 

migrationes sistens, &>c. : 526. Ley den. 




PRESENTED 

- 1 APk WW 



PLATE 3 

Figs, i and 2. Pencil sketches by Sydney Parkinson of kangaroos seen 
at Endeavour River, Queensland, in July 1770. (Preserved in the 
Zoological Library of the British Museum (Natural History)) 



B.M. (N.H.), Zoology I, 3 



PLATE 3 




Fig. i 






\ 




.. 




Fig. 2 



PLATE 4 

Fig. 3. Wash drawing by Nathaniel Dance of the skull of a young 
kangaroo (Macropus robustus subsp.) obtained at Endeavour River, 
Queensland, in July 1770 

Fig. 4. Lower jaw of the skull in Fig. 3 




B.M. {N.H.), Zoology I, 3 



PLATE 4 




™» 2 ! 1 1 1 6 I 



ijo l|l ' 1J3 tjj jft 



Fig. 3 





•1 5; 6! 






I H ^'J „ 



Fig. 4 



PLATE 5 

Fig. 5. Photograph of the skull of a young Great Grey Kangaroo 
obtained at Endeavour River, Queensland, by Captain Cook's party 
in July 1770. This plate is the photo-lectotype of Macropus canguru 
(Miiller). The specimen, which no longer exists, was number 1732 
in Owen's Catalogue (1853) and number 3703 in Flower's Catalogue 
(1884). Scale unknown 




Bull. B.M. (N.H.), Zoology I, 3 



PLATE 5 




Fig. 5 






»"«® 






PRESENTED 

- 1 AFk 1950 



PRINTED IN 

GREAT BRITAIN 

AT THE 

UNIVERSITY PRESS 

OXFORD 

BY 

CHARLES BATEY 

PRINTER 

TO THE 
UNIVERSITY 




<5 SEP 1950 

NOTES ON ASTEROIDS IN THE BRITISH 
MUSEUM (NATURAL HISTORY) 

D. DILWYN JOHN 



LERNAEODISCUS PUSILLUS NOV. SPEC. 

A RHIZOCEPHALAN PARASITE OF A 

PORCELLANA FROM EGYPT 

H. BOSCHMA 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ZOOLOGY Vol. i No. 4 

LONDON : 1950 



NOTES ON ASTEROIDS IN THE 

BRITISH MUSEUM 

(NATURAL HISTORY) 



BY 



D. DILWYN JOHN 



LERNAEODISCUS PUSILLUS NOV. SPEC. 

A RHIZOCEPHALAN PARASITE OF 

A PORCELLANA FROM EGYPT 

BY 

DR. HILBRAND BOSCHMA 




Pp. 51-65; Pi. 6; 4 Text-figures 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ZOOLOGY Vol. 1 No. 4 

LONDON: 1950 



THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted in 1949, is to be 
issued in five series, corresponding to the Departments 
of the Museum. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four hundred 
pages, and will not necessarily be completed within one 
calendar year. 

These papers form Vol. I, No. 4, of the Zoological 
series. 



PRINTED BY ORDER OF THE TRUSTEES OF 
THE BRITISH MUSEUM 

Issued August 1950 Price Four Shillings 



NOTES ON ASTEROIDS IN THE BRITISH 
MUSEUM (NATURAL HISTORY) 

2. SOME ASTROPECTINID SPECIES 

By D. DILWYN JOHN 

(director of the national museum of wales, Cardiff) 

(With Plate 6) 

The first Note in this series (John, 1948) began with the statement that the Asteroids 
in the British Museum (Natural History) were being revised. This, the second Note, 
will be the last in the series by the present author, who has since left the Museum 
stall. It is shorter than it was intended to be and deals only with the following six 
Astropectinid species: 

Lonchotaster tartareus Sladen. Leptychaster antarcticus Sladen. 

Dytaster exilis Sladen. Leptychaster kerguelensis Smith. 

Plutonaster agassizii (Verrill). Craspidaster hesperus (Muller & Troschel). 

Lonchotaster tartareus Sladen 

Lonchotaster tartareus Sladen, 1889, Rep. Voyage Challenger (Zool.), 30 : 104, pi. 16, figs. 1-5. 

The only species and the only specimens of the genus Lonchotaster remain those 
described by Sladen in 1889, L. tartareus from 2,400 fathoms between the Canaries 
and the Cape Verde Islands, and L. forcipifer from nearly 2,000 fathoms in the 
Southern and Antarctic Oceans south-west of Australia. The large Astropectinid 
described by H. L. Clark (1916: 30) as Lonchotaster magnificus was referred to Dipsa- 
caster by Fisher (19 19: 150). 

Fisher, both in 1917 (p. 170) and 1919 (p. 150), makes what are, in effect, minor 
corrections to Sladen's account of L. tartareus, saying there is a small spine on each 
marginal plate and one on most of the actinal intermediate plates; he refers to 
Sladen's figures as bearing out his statement. As for the superomarginal plates, 
Fisher is wrong and Sladen's account, with which his plate agrees, is correct: 'within 
the interbrachial arc and at the base of the rays in the large example, a small conical 
tubercle is present close to the upper end of the plate, but it is not found in the 
smaller specimens'. For the inferomarginals neither Sladen's account nor Fisher's is 
quite correct. In the larger specimens there are small spines, of diminishing size, as 
far out as about the thirtieth plate, but not beyond ; they are present on the plates 
of the interbrachial arc of one of the smaller, entirely absent from the other. 

Sladen's account of the spination of the actinal intermediate plates is correct, 
including the implication that there are no spines on those of the smaller specimens. 



54 NOTES ON ASTEROIDS IN THE BRITISH MUSEUM (NAT. HIST.) 

Dytaster exilis Sladen 

Dytaster exilis Sladen, 1889, Rep. Voyage Challenger (Zool.), 30 : 65, pi. 2, figs. 3 & 4 ; pi. 4, figs. 9 
& 10 (figs, of var. gracilis) ; Wood-Mason & Alcock, 1891: 429; Alcock, 1893: 80. 

The Challenger took the type of D. exilis off Valparaiso in the Pacific, those of its 
varieties gracilis and carinata in the Atlantic near Tristan da Cunha and off the 
Maryland coast of N. America respectively. The only subsequent records are those 
of exilis itself by Wood-Mason and Alcock from the Bay of Bengal, where it 'has 
several times been met with . . . between 1748 and 1924 fathoms on globigerina ooze \ 
They did not describe their specimens beyond giving the colour when fresh as salmon- 
pink. 

One of their specimens, from St. 117, 1,748 fms., is in the British Museum. It is 
dry and small: R = 47 mm., r = 9 mm., R : r is 5-2. The abactinal paxillae have four 
to ten finely thorny spinelets ; there are no pedicellariae among them. The supero- 
marginals number thirty- three. They are not confined to the lateral wall but encroach 
a little on the abactinal surface ; those in the inter-brachial angle do so to the extent 
of 1 mm. This is a marked difference to the type of exilis ; in the variety gracilis, on 
the other hand, they do encroach abactinally though not so strongly as in this 
specimen. When seen from the side the length of the plates is less than the height in 
the inter-brachial angle, greater than it in mid-arm, equal to it at the end of the arm. 
The large spines are missing from the plates at the ends of the arms which are 
abraded, but I am unable to say if they have merely been rubbed off. 

The inferomarginals correspond to and are of the same size as the superomarginals 
as seen from the side. On the actinal surface their breadth is greater than their length 
on the inner part of the ray. In the interbrachial angle some of the marginal plates 
of both series carry two spines. 

The enlarged spine on the adambulacral plate first appears about half-way down 
the arm and arises more often from the second than the first comb of spines. The 
latter has ten, the former eight, spines, and they are followed by a third row as 
Sladen describes for exilis. The actinal intermediate plates extend to about the third 
inferomarginal. Each bears a group of widely spaced spines, up to fourteen on the 
largest. They and the spines of the marginal and adambulacral plates are finely 
thorny. 

The madreporite is neither large nor conspicuous. 

In the shape of the superomarginal plates, the absence of pedicellariae, and the 
occurrence of the enlarged spine on the adambulacral plates I see this specimen as 
nearer to the var. gracilis than to exilis itself. Experience with other species leads me 
to believe it possible that more specimens may serve to bridge the gap which now 
appears to exist. 

Verrill (1895 : 131) was not able to satisfy himself that D. exilis var. carinata was 
distinct from the young of his D. grandis (of which D. madreporifer Sladen is a 
synonym). A direct comparison leaves no doubt of its distinctness. In the first place 
the larger specimen described by Sladen cannot be regarded as young, having R = 
98 mm. The paxillae of its disk are comparatively large, those of grandis conspicuously 
small ; the pedicellariae on the actinal intermediate plates of carinata are larger and 



NOTES ON ASTEROIDS IN THE BRITISH MUSEUM (NAT. HIST.) 55 

of valves more highly modified than those of grandis (Plate 6, fig. 1) ; the adambulacral 
armature differs, for whereas grandis has only one row of strong furrow spines, 
carinata has two, the second being of the peculiar dagger-like form described by 
Sladen. Finally, the appearance of the two forms is quite different to the naked eye 
for, whereas D. grandis is distinguished by the strong high sides which the marginals 
give to its rays, in the var. carinata the marginals are comparatively poorly developed, 
their combined height being only a little more than half that of grandis, and the spines 
are correspondingly smaller (Plate 6, figs. 2 & 3). 

Plutonaster agassizii (Verrill) 

Archaster agassizii Verrill, 1880, Amer. J. Sci. 20 : 403. 

Plutonaster rigidus Sladen, 1889, Rep. Voyage Challenger (Zool.), 30 : 91, pi. 14, figs. 3 & 4 ; pi. 

15, figs. 3 & 4; Koehler, 1909: 19, pi. 4, fig. 6; pi. 10, figs. 5 & 6. 
Plutonaster rigidus var. semiarmata Sladen, 1889, Rep. Voyage Challenger {Zool.), 30 : 94, pi. 14, 

fig- 5- 
Plutonaster agassizii Verrill, 1894, Proc. U.S. Nat. Mus. 17 : 248; 1895: 131 ; 1899: 211, pi. 27, 
fig. 6. 

Verrill (1880 : 403) in his ' Notice of the remarkable Marine Fauna occupying the 
outer banks off the Southern Coast of New England' described the new species 
Archaster agassizii. Sladen (1889) made no reference to Verrill's paper in the Chal- 
lenger Report. In 1894 (p. 248) Verrill placed his species in Sladen's genus Plutonaster ; 
listed Sladen's rigidus and rigidus var. semiarmata and a part of his bifrons, all from 
off the coast of North America, as synonyms ; and added to the description. In 1899 
he described the species as occasionally having pedicellariae and gave a figure show- 
ing one. 

Koehler (1909: 19) used Sladen's name, rigidus, for describing a series taken in 
mid- Atlantic in the latitude of the Azores, explaining that he did so because he 
could not be sure that Verrill's agassizii and Sladen's rigidus were the same. He 
found Verrill's description inadequate and his attempt to have photographs of his 
specimens compared with Verrill's had failed. 

Dr. Austin Hobart Clark has generously made it possible for me to make the sort 
of comparison that Koehler wished to make by sending me six specimens of Verrill's 
species. They came from off New Jersey, 39 58' 30* N., 70 30' 00" W., 384 fms. 

They show that agassizii and rigidus are one. Koehler had found that the var. 
semiarmata of Sladen could not be maintained, so variable is the occurrence of spines 
on the inferomarginal plates. Verrill (1894: 248) says that there may be all grada- 
tions from those having no marginal spines whatever to those that have a large spine 
on nearly every marginal plate of both series. Koehler does not record spines on the 
superomarginal plates and it may be assumed that they were not present in his 
specimens. There is none in the six specimens from Verrill before me, but in the 
type of Sladen's rigidus there is on one or two plates a single slightly enlarged granule 
such as I have seen to occupy a similar position from which a spine often arises in 
other asteroids. 

Koehler makes no mention of pedicellariae. I find a row of four to be present 
actinally in the midline of one interradius of one of Verrill's specimens, and a single 



56 NOTES ON ASTEROIDS IN THE BRITISH MUSEUM (NAT. HIST.) 

one in another interradius. They have four or five blades. The type of rigidus has 
some small groups of spines in the actinal intermediate areas which are pedicellaria- 
like in their disposition, but the ' blades ' are short and coarse. 

Sladen (p. 92) described the conical spinelet immediately behind the furrow spines 
on the outer adambulacral plates. Though Koehler did not mention it, it is to be 
assumed it was present since he identified his specimens with Sladen's species. It is 
present in VerriU's specimens, more strongly developed in some than in others. 

R : r is more than 3 in one of VerriU's specimens (R = 49 mm., r = 15 mm.) ; it is 
less than 3 in the remaining five in which R varies from 42 to 63 mm. and r from 
17 to 22 mm. 

Verrill included the small specimen which Sladen (p. 88) described with a query as 
P. bifrons in his synonymy of agassizii. It possesses a spine on each marginal plate, 
inferior and superior ; there is a large spine behind the furrow series on each adambula- 
cral plate. In view of its origin it is probably the young of agassizii, but it cannot be 
said with certainty that it is. 1 

Leptychaster antarcticus Sladen and L. kerguelensis Smith 

Lepty chaster antarcticus Sladen, 1889, Rep. Voyage Challenger (Zool.), 30 : 190, pi. 31, figs. 3 & 4; 

pi. 32, figs. 7 & 8. 
Leptychaster kerguelensis Smith, 1876, Ann. Mag. Nat. Hist. 17 : no. 

The type of L. antarcticus, and a second and smaller specimen taken with it (R = 
10-5 mm., r = 4-5 mm.), are in the Museum collection. They are the only specimens 
recorded. Bell (1908 : 9) thought them the young of kerguelensis, but he gave no good 
reasons for doing so. 

Koehler (1917: 53) discussed the question and Fisher (1940: 83) referred to it, but, 
while not affirming that Bell was wrong, neither accepted his conclusion. It seemed 
well that I, with access to the types of both species, should re-examine them and 
other available specimens and report what I find. 

The paxillae of the greater part of the swollen abactinal surface of the type of 
antarcticus have lost their spines. It may have happened during transport to and from 
a safe place in the Second World War. They appear to have been present when the 
Challenger figure (pi. 31, fig. 3) was made. While Sladen's written description is of 
his usual excellence, fig. 4, pi. 31, is a poor representation: it is, indeed, a misrepre- 
sentation of the mouth plates, which are as Sladen describes them in words. It is 
hoped that the photograph given here conveys a better idea (Plate 6, fig. 4). 

Sladen's description of kerguelensis is of a large specimen of R = 66 mm. ; though 
he listed smaller specimens and gave their sizes he did not otherwise describe them. 
He states (p. 192) that kerguelensis is distinguished from antarcticus by the longer and 
more cylindrically rounded rays, by the larger and more compact paxillae, by the 
smaller actinal intermediate areas, and, above all, by the characteristic adambulacral 
armature. 

The smallest specimen of kerguelensis in the collection was taken with three larger 

1 A doubt is possible about its origin. On p. 87 Sladen gives it as St. 47, off the coast of N. America. 
On p. 88 he gives St. 47a. There was no Challenger station of that number but there was one by the 
Porcupine and it was in the Faroe Channel. 



NOTES ON ASTEROIDS IN THE BRITISH MUSEUM (NAT. HIST.) 57 

specimens (R up to 60 mm.) in 50 fms., off Marion Is. In it R == 13-8 mm. and r = 5 mm., 
so that it is slightly smaller than the type of antarcticus (R = 15 mm., r = 6 mm.). 
A direct comparison has been made between them. The rays of the kerguelensis 
specimen are, in proportion, longer and more rounded, and the actinal intermediate 
areas are smaller; and the differences in proportion give a different facies to each 
specimen. 

But the paxillae are similar in the two specimens and as Sladen described them 
for antarcticus, though his figure is not very good. It is, however, far better than is 
that of the paxillae of kerguelensis (pi. 32, fig. 1). In only three of the fifteen Museum 
specimens are they as shown in that figure, with the spines represented by low 
rounded granules, tending to be polygonal where crowded. In the others they are 
much more spine-like and radiate apart. Though it is not necessarily the biggest 
specimens in which the paxillae spines are lowest and most crowded, it is in the 
smallest that they are most spine-like. In short, the distinction between kerguelensis 
and antarcticus based upon the nature of their paxillae appears not to be real. 

The question of the adambulacral armature remains. It can only be said that 
Sladen's descriptions are correct and that his figs. 2 & 8, pi. 32, are good representa- 
tions. It may be added that Koehler's eight specimens of kerguelensis conformed 
with Sladen's description for that species, and that it is implicit in Fisher's account 
that his three specimens also did so. 

And so, since no intermediate stages have been found, it seems best to go on regard- 
ing kerguelensis and antarcticus as distinct species distinguished by their different 
adambulacral armature. 

The three starfishes from the Cape which Bell (1905 : 242) recorded as L. kerguelensis 
are Dipsacaster sladeni Alcock, as Mortensen (1933: 237) pointed out. Bell (1908: 9) 
also recorded the species from the Ross Sea, including one specimen in which R == 
212 mm. I cannot find that specimen ; nor are there any Ross Sea specimens labelled 
L. kerguelensis. There are several jars labelled by Bell ' Lepty chaster young' or 'very 
young', and I suppose them to be the young examples to which he referred. They 
are, however, not Leptychaster but Odontaster — and some other genera are included. 

Craspidaster Hesperus (Miiller & Troschel) 

Archaster hesperus Miiller & Troschel, 1840, Ber. preuss. akad. Wiss. : 104. 

Craspidaster hesperus Sladen, 1889, Rep. Voyage Challenger (Zool.), 30: 177, pi. 17, figs. 5-7; 

pi. 18, figs. 1-4; Doderlein, 1921: 5 (for synonymy), 8, pi. 1, figs. 2-3. 
Craspidaster glauconotus Bedford, 1900, Proc. Zool. Soc. Lond.: 290, pi. 24, figs. 8a, b; Doderlein, 

1 921: 8, pi. 1, figs. 4-6. 
Craspidaster hesperus crassus Doderlein, 192 1, Siboga Exped. Monog. 46 i : 9, pi. 1, figs. 1 & la. 

There are in the British Museum thirty-nine specimens. One is from an unknown 
locality, five are said to be from Japan but there can be no certainty of it, twenty-one 
from the Chusan Archipelago, one from Amoy, and another from Hong Kong (Chal- 
lenger), two each from the Philippines (Challenger) and Batavia, and six specimens of 
Bedford's glauconotus from Malacca. 

Doderlein had twelve specimens and took into account, for measurements, &c, 
three more. He recognized three sub-species differing from one another in the length 



58 



NOTES ON ASTEROIDS IN THE BRITISH MUSEUM (NAT. HIST.) 



and width of the arm, the number, size, and spination of the marginal plates, and the 
number and nature of the actinal intermediate plates. Four of his specimens were 
from China and Japan, the remainder from East Indian or Malayan seas. The former 
had shorter and wider arms, and larger and — on the whole, and especially in the 
second row — fewer actinal intermediate plates. One of the Chinese specimens of un- 
usually plump form, with massive marginals and having only one row of actinal inter- 
mediate plates, he made the type of a new sub-species, crassus; the remainder he 
regarded as typical hesperus. The Malayan examples, with longer more slender arms, 
more numerous marginals, smaller and more actinal intermediate plates — especially 
in the second row — and with, in the larger, spines on the ventral faces of the infero- 
marginals, he grouped with Bedford's specimens in the sub-species glauconotus. 

The present collection bears out Doderlein's conclusions concerning the relation of 
R : r, and the number of marginal plates. In the twenty-one Chusan specimens R 
ranges from 8-5 to 42 mm. and the relation R : r varies from 2-1 in the smaller to 3-5 
in the larger. In the six specimens of glauconotus from Malacca the range of R is 18 
to 67 mm. and oiR'.r 3-2 to 4-6. There is no doubt that the latter are conspicuously 
longer-armed. They have, too, a larger number of superomarginal plates. Perhaps 
the most telling way of making a difficult comparison is to bring together (1) a number 
of specimens of roughly equal sizes, as follows : 



Locality 


R in mm. 


R:r 


No. of marginals 


? Japan .... 
Chusan .... 
Timor (Doderlein) 
Malacca {glauconotus) 


34 

29-5 

29 

31 


3-2 
3-i 

3-6 
4'4 


24 
23 
26 

33 



and (2) a number of specimens with roughly equal numbers of marginal plates 



Locality 


No. of marginals 


R 


R :r 


? Japan .... 


27 


4 1 


3-4 


Chusan .... 


30 


42 


3-5 


Hong Kong 


3i 


53 


3-6 


Philippines .... 


3i 


37-5 


3-8 


Malacca (glauconotus) 


33 


3i 


4-4 



The first list shows that Bedford's glauconotus is sharply marked off from the other 
specimens by the high value of R : r and by the large number of marginal plates ; the 
second, that a specimen of glauconotus with a given number of marginals is of much 
smaller major radius and has a markedly higher value of R : r than specimens of 
hesperus with the same number of marginals. 1 Each list tells the same story, but by 
means of different specimens. 

One of the Batavia specimens is roughly equal in size (R = 57 mm.) to one of those 
from Malacca (R = 59 mm.). R : r is 4 in the former, 4-3 in the latter, and the relative 
numbers of marginal plates are 40 and 47. 



1 The large major radius of the Hong Kong (Challenger) specimen is because of its peculiarly massive 
marginals; compare the type of crassus which, with only 20-22 marginals, has R = 46 mm. 



NOTES ON ASTEROIDS IN THE BRITISH MUSEUM (NAT. HIST.) 59 

The spines on the lower surfaces of the inferomarginal plates and on the actinal 
intermediate plates afford a strong difference between Bedford's glauconotus and 
typical hesperus. They are well developed on each of the six specimens. They occur, 
strongly on the inferomarginal plates, poorly developed on the actinal intermediate 
plates, of the larger specimen (R = 57 mm.) from Batavia ; there are traces of them 
on the actinal-intermediate plates only of the second Batavian specimen (R = 
57 mm.). There are spines, varying in number but never numerous, on the lower 
surfaces of the inferomarginals of (1) the Challenger specimen from Hong Kong (an 
odd one or two), (2) the larger Challenger specimen from the Philippines (one on each 
of two rays) , and (3) one of the Japan specimens (one on each of the first eight plates) . 

I find nothing to support Doderlein's implication that there is a real difference in 
the number of actinal intermediate plates of 'Chinese' and 'Malayan' specimens. 
He gives as a characteristic of some of the former that they have few and massive 
plates, sometimes only one row (var. crassus). It is true that in the British Museum 
collection six of the smaller specimens from Chusan (R = 10-17 mm.) have only one 
row, but since the remaining and larger specimens have two rows, and the largest 
specimens have the highest number of plates, this is clearly a matter of growth. The 
only other specimens with no second row of actinal intermediate plates are (1) one 
of glauconotus of no less than R = 60 mm. (no second row in two interradii ; a single 
plate comprises the ' second row ' in each of the other three) ; (2) the smallest specimen 
of glauconotus (R = 18 mm.) ; (3) Sladen's 'young phase' (R = 22 mm.) from the 
Philippine Islands. The largest glauconotus (R = 67 mm.) has six to eight plates in 
the first, three plates in the second, row. The specimen from an unknown locality is 
exceptional: it has R = only 31 mm. and yet has seven to eight plates in the first row, 
three to four in the second, and it possesses a third row of one plate on either side. 

Sladen described the occurrence of a thumb-like spine on the aboral margin of the 
adambulacral plates of his Hong Kong specimen and its absence from those from the 
Philippines. It was not present in the specimens from the Philippines seen by Fisher 
(1919: 60). Doderlein does not mention it. 1 It is (as Bedford says) present in glauco- 
notus ; I find it in each specimen from the smallest (R = 18 mm.) to the largest 
(R = 67 mm.). It is present in the specimen from an unknown locality and in that 
from Amoy, in three of those from Japan (R = 35-41 mm.), but it is absent from all 
but a few plates of the fourth (R = 34 mm.). It is not present in the two specimens 
from Batavia. It is absent from twenty of the twenty-one specimens from Chusan 
of R = 8-5 to 29-5 mm., but is present in the twenty-first which is conspicuously 
larger having R = 42 mm. 

The conclusion appears to be that in the present state of our knowledge glauco- 
notus should continue to rank as a sub-species distinguished by the length of its rays, 
the number of its marginals, and the presence of spines on the inferomarginal and 
actinal intermediate plates ; but that crassus cannot be maintained. The species is 
seen to be variable: e.g. the Hong Kong specimen approaches Doderlein's crassus in 
its massive marginals and yet bears traces of spines, a glauconotus character, on some 
of them ; the thumb-like spine of the adambulacral plate is absent from most small 

1 His fig. 6a on pi. i shows it to have been absent from his specimen from Lombok. Text-fig. i and the 
accompanying text do not make clear the possibility of its existence. 



60 NOTES ON ASTEROIDS IN THE BRITISH MUSEUM (NAT. HIST.) 

specimens but it is present in one glauconotus, R = 18 mm., and it may be entirely 
wanting on large specimens up to R = 57 mm. 

REFERENCES 

Alcock, A. 1893. An account of the collection of deep-sea Asteroidea. Ann. Mag. Nat. Hist. 

11 : 73-121, 3 pis. 
Bedford, F. P. 1900. On Echinoderms from Singapore and Malacca. Proc. Zool. Soc. Lond.: 

271-299, 4 pis. 
Bell, F. J. 1905. The Echinoderma found off the coast of South Africa. 2. Asteroidea. Mar. 

Invest. S. Afr. 3 : 241-253. 
1908. Echinoderma. National Antarctic Expedition, igoi-igo^, Natural History, Zoology 

(Echinoderma), 4 : 1-16, 5 pis. 
Clark, H. L. 19 16. Report on the Sea-Lilies, Starfishes, Brittlestars and Sea-Urchins obtained 

by the F.I.S. Endeavour on the coasts of Queensland, New South Wales, Tasmania, Victoria, 

South Australia, and Western Australia. Biol. Res. 'Endeavour', igog-igi4, 4: 1-123, 

44 pis. 
Doderlein, L. 192 1. Die Asteriden der Siboga Expedition. I. Porcellanasteridae, Astro- 

pectinidae, Benthopectinidae. Siboga Exped. Monog. 46 i : 1-47, 13 pis. 
Fisher, W. K. 1917. Notes on Asteroidea. Ann. Mag. Nat. Hist. 20 : 166-172. 
1919. Starfishes of the Philippine Seas and adjacent waters. Bull. U.S. Nat. Mus. 100 

(3): 1-712, 156 pis. 

1940. Asteroidea. 'Discovery' Rep. 20 : 69-306, 23 pis. 

John, D. D. 1948. Notes on Asteroids in the British Museum (Natural History) — 1. The Species 

of Astropecten. Novit. Zool. 42 : 485-508, 4 pis. 
Koehler, R. 1909. Iichinodermes provenant des campagnes du yacht Princesse Alice. Result. 

Camp. sci. Monaco, 34 : 1-3 17, 32 pis. 
1917. fichinodermes recueilles par M. Rollier du Baty aux lies de Kerguelen, en 1913-1914. 

Ann. Inst. Oceanogr. Monaco, 7 (8) : 87 pp., 10 pis. 
Mortensen, T. 1933. Echinoderms of South Africa. Vidensk. Medd. naturh. Foren. Kbh. 93 : 

215-400, pis. 8-19. 
Muller, J., & Troschel, F. H. 1840. [In Report of the Session of 30 April 1840.] Ber. preuss. 

akad. Wiss. 
Sladen, W. P. 1889. Asteroidea. Rep. Voyage Challenger (Zool.), 30. 
Verrill, A. E. 1880. Notice of the remarkable Marine Fauna occupying the outer banks off the 

southern coast of New England. Amer. J. Sci. 20 : 390-403. 
1894. Description of new species of Starfishes and Ophiurans, with a revision of certain 

species formerly described. Proc. U.S. Nat. Mus. 17 : 245-297. 
1895. Distribution of the Echinoderms of North-eastern America. Amer. J. Sci. 49: 

127-141. 
1899. Revision of certain genera and species of Starfishes with descriptions of new forms. 

Trans. Conn. Acad. Arts Sci. 10 : 145-234, 8 pis. 
Wood-Mason, J., & Alcock, A. 1891. Natural History notes from H.M. Indian Marine Survey 

Steamer Investigator. Ser. 2, no. 1. On the results of deep-sea dredging during the season 

1 890-1891. Ann. Mag. Nat. Hist. 8 : 427-443. 



PLATE 6 

Fig. i. Dy taster exilis var. carinata, type, mouth-angle and actinal- 
intermediate area, X5. 

Fig. 2. Dy taster exilis var. carinata, type, side view of the proximal 
portion of arm, x 4. 

Fig. 3. Dytaster grandis, cotype, side view of proximal portion of arm, 
X 4 . 
Fig. 4. Lepty chaster antarcticus, type, under surface of disk, x 10. 



B.M. {N.H.) Zoology I, 4 



PLATE 6 




LERNAEODISCUS PUSILLUS NOV. SPEC, 

A RHIZOCEPHALAN PARASITE OF A 

PORCELLANA FROM EGYPT 



By HILBRAND BOSCHMA 

(director, rijksmuseum van natuurlijke historie, leiden) 

In 1936 Dr. Isabella Gordon kindly sent me twelve specimens of Rhizocephalan 
parasites on Porcelain Crabs collected by Dr. R. Gurney in coral rock on the Harbour 
Reef near Ghardaqa, Red Sea, Egypt. The hosts of these parasites were provisionally 
identified as Porcellana serratifrons of Nobile, nee Stimpson. The parasites appear to 
represent a hitherto undescribed species. 




Fig. 1. Lernaeodiscus pusillus: a-c, dorsal view of three specimens, mantle opening in the upper 
part, stalk in the lower part of the figures ; d-f, ventral view of the same specimens, x 18. 

The animals are of very small size, their greatest diameter being about 2 mm., 
their antero-posterior diameter (in the median plane) about i^ mm., and their smallest 
(dorso-ventral) diameter less than 1 mm. The total diameter in the antero-posterior 
direction is, as a rule, slightly less than the greatest diameter. The outlines of three 
specimens in dorsal view are given in Fig. xa-c, in ventral view in Fig. id-f. The 
shape of the parasites is more or less roundish or somewhat trapezoid or triangular ; 
their contour is slightly irregular as the mantle shows a number of rather incon- 
spicuous lappets. The comparatively wide mantle opening, which is surrounded by a 
well-developed muscular wall, is found on the anterior region of the dorsal surface. 
As a rule the dorsal surface shows a system of three shallow grooves running from 

zool. 1. 4 h 




Fig. 2. Lemaeodiscus pusillus, specimen of Fig. la, d. Transverse section showing one of the 
colleteric glands {eg) . dms, dorsal mesentery ; mc, mantle cavity ; v m, visceral mass ; vms, ventral 
mesentery, x 60. 




Fig. 3. Lemaeodiscus pusillus, specimen of Fig. la, d. Central parts of transverse sections, a from 
a region not far from the stalk, each following section from a more anterior region, dms, dorsal 
mesentery ; //, left testis ; Ivd, left vas deferens ; mc, mantle cavity ; rt, right testis ; rvd, right vas 
deferens ; st, stalk ; vm, visceral mass ; vms, ventral mesentery, x 64. 



LERNAEODISCUS PUSILLUS NOV. SPEC. 63 

the centre to the mantle opening and to the lateral parts of the posterior region of the 
body. On the ventral surface there is a distinct groove running from the stalk in an 
anterior direction ; this groove varies in length and in breadth. 

The three specimens shown in Fig. 1 were sectioned transversely for the study of 
their internal structure. In sections from the region about half-way between the 
stalk and the mantle opening the colleteric glands are found ; as a rule one of these is 
situated more anteriorly than the other. These glands (Fig. 2, eg) are more or less 
cup-shaped small cavities surrounded by an epithelium with a stronger affinity for 
stains than the surrounding parts. The figure further shows that the dorsal surface 
of the visceral mass is broadly attached to the mantle, in this way forming the so- 
called dorsal mesentery. On the other side the visceral mass is connected with the 
mantle by means of a real mesentery, the ventral mesentery. Where the latter is 
attached to the mantle there is, externally, the longitudinal groove referred to above. 

In the three sectioned specimens the colleteric glands entirely agree with one 
another in shape, their position in the visceral mass, and their size. The male organs 
in two of the sectioned specimens are also similar in every respect (Fig. 3), but in the 
third specimen (Fig. 4) they are slightly more complicated. 

The male organs closely correspond with those of Lernaeodiscus okadai Boschma 
(cf. van Baal, 1937, figs. 18-21). The male openings, in a region about half-way 
between the stalk and the mantle opening, are found on each side of the ventral 
mesentery (Fig. 4^, e). The vasa deferentia run along the ventral mesentery until 
they reach the posterior part of the visceral mass. Here they turn towards the dorsal 
surface (Figs. 3a, 4a), and continue their course along the dorsal mesentery in an 
anterior direction. After the vasa deferentia have passed into the testes the latter 
extend in a lateral direction, so that the terminal part of the testes is the most lateral 
part of the male organs (Fig. 36, c). 

As remarked above, the male organs in two of the sectioned specimens have a 
similar shape (as represented in Fig. 3) ; in the third specimen the male organs show 
some differences. Here the left testis (Fig. 4^, e) does not extend in a lateral direction, 
whilst the terminal part of the right testis after continuing its course in a lateral 
direction towards the right margin of the visceral mass {a in Fig. 4) obtains a curved 
shape by extending towards the median plane again (p in Fig. 4) . The closed end of 
this testis consequently lies next to the right vas deferens (Fig. 46). 

Besides having a course in a lateral direction the testes in all the three specimens 
are strongly contorted, so that in sections they appear to be divided into numerous 
smaller parts. 

It is rather difficult to define the characters by which Lernaeodiscus pusillus can 
be distinguished from the other species of the genus that are, like the new species, 
parasites of Porcelain Crabs, viz. L. porcellanae Miiller (cf . Muller, 1862 ; Boschma, 
1931) and L. okadai Boschma (cf. Boschma, 1935 ; van Baal, 1937). 

The external shape of Lernaeodiscus porcellanae seems to be rather constant, the 
animal having well-developed lappet-like expansions of the mantle. But too few 
specimens are known to establish this peculiarity as a constant character for full- 
grown as well as immature specimens. In L. okadai, van Baal (1937) has shown that 
the external shape is subject to a very large amount of variation. Here, as a rule, 





Fig. 4. Lemaeodiscus pusillus, specimen of Fig. ic, /. Central parts of transverse sections, a from 
a region not far from the stalk, each following section from a more anterior region, a, anterior 
part of right testis; dms, dorsal mesentery; It, left testis; Ivd, left vas deferens; mc, mantle 
cavity ; p, posterior part of right testis ; ro, right male genital opening ; rvd, right vas deferens ; 
vm, visceral mass ; vms, ventral mesentery, x 64. 



LERNAEODISCUS PUSILLUS NOV. SPEC. 65 

the lappets do not occur in young specimens but are generally distinct in mature 
animals. The specimens of L. pusillus have, as far as their external shape is concerned, 
a rathei constant appearance. 

The colleteric glands in the genus Lernaeodiscus are of such a simple structure that 
they cannot furnish characters for specific distinction. 

The male genital organs are, to a large degree, subject to individual variation, as 
is evident from van Baal's (1937) elaborate researches on numerous specimens of 
L. okadai. 

The only remaining distinctive character is that of the size of the animals. On this 
character L. porcellanae, by its comparatively large size, is at once distinguished from 
>L. okadai and L. pusillus. In L. pusillus the greatest diameter is about 2 mm., and 
the total length is but slightly smaller. The sectioned specimens are fully mature, as 
their mantle cavities contain large quantities of eggs. For L. okadai there are the 
following data (the numbers giving the length and the greatest transverse diameter 
in mm.) recorded by van Baal (1937) : 

2JX3 (small number of eggs) ; 4x5^ (no eggs) ; 4^x5 (small number of eggs) ; 
4x5 (large number of eggs) ; 2f X3J (very small number of eggs) ; lJX2 (no 
eggs); 6X7J (large number of eggs); 3iX5i (no eggs); 2X4J (many eggs); 
2|X4 (crowded with eggs); 5jx6 (many eggs); 4^x6 (many eggs); 4x4^ 
(many eggs) ; 2 J X 4 (without eggs) . 

These data show that the specimens with numerous eggs are the larger ones in 
which at least one dimension reaches 4 mm. Moreover, when in large specimens no 
eggs are present in the mantle cavity they may have been recently discharged from 
this cavity. The data, therefore, give sufficient evidence for the opinion that L. 
okadai reaches its mature state at a stage in which at least in one dimension the body 
has a size of 4 mm. On the other hand, L. pusillus is fully mature at a size of 2 mm. 

Summarizing it may be remarked that though the specific characters of Lernaeo- 
discus pusillus may appear unconvincing there is sufficient evidence for regarding 
the parasite as specifically distinct from the other forms belonging to the genus. 

REFERENCES 

Baal, I. van. 1937. Biological Results of the Snellius Expedition. II. Rhizocephala of the 
Families Peltogastridae and Lernaeodiscidae. Temminckia, 2 : 1-94, 3 pis. 

Boschma, H. 1931. Papers from Dr. Th. Mortensen's Pacific Expedition, 1914-1916. LV. Rhizo- 
cephala. Vidensk. Medd. naturh. Foren., Kbh. 89 : 273-380. 
— 1935. Notes on Japanese Rhizocephala, with Description of two new Species. Zool. Meded. 
18 : 1 51-160. 

Muller, F. 1862. Die Rhizocephalen, eine neue Gruppe schmarotzender Kruster. Arch. 
Naturgesch. 28 : Bd. 1 : 1-9. 




PRESENTED 

5SE1 ibjw 



PRINTED IN 

GREAT BRITAIN 

AT THE 

UNIVERSITY PRESS 

OXFORD 

BY 

CHARLES BATEY 

PRINTER 

TO THE 
UNIVERSITY 




9 - NOV 1951 

ON A^&s&KE DEEP-SEA FISH 
NOTACANTHUS PHASGANORUS 

GOODE 

(HETEROMI-NOTACANTHIDAE) 

FROM THE ARCTIC BEAR ISLE 

FISHING-GROUNDS 

DENYS W. TUCKER and J. W.JONES 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ZOOLOGY Vol. i No. 5 

LONDON :i9 5 i 



ON A RARE DEEP-SEA FISH 

NOTACANTHUS PHASGANORUS GOODE 

(HETEROMI-NOTACANTHIDAE) 

FROM THE ARCTIC 
BEAR ISLE FISHING-GROUNDS 

BY 

DENYS W. TUCKER , B.Sc. 

AND 

J. W.JONES, Ph.D. 
Pp. 67-79; JPk- 7-9 




BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ZOOLOGY Vol. 1 No. 5 

LONDON : 1951 



THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted in 1949, is issued in 
five series, corresponding to the Departments of the Museum. 

Parts appear at irregular intervals as they become ready. 
Volumes will contain about three or four hundred pages, 
and will not necessarily be completed within one calendar 
year. 

This paper is Vol. 1, No. 5, of the Zoology series. 



PRINTED BY ORDER OF THE TRUSTEES OF 
THE BRITISH MUSEUM 

Issued November 1951 Price Five shillings 



ON A RARE DEEP-SEA FISH 

NOTACANTHUS PHASGANORUS GOODE 

(HETEROMI-NOTACANTHIDAE) FROM THE ARCTIC 

BEAR ISLE FISHING-GROUNDS 

By DENYS W. TUCKER, B.Sc. 

(BRITISH MUSEUM (NATURAL HISTORY)) 

and 

J. W. JONES, Ph.D. 

(UNIVERSITY OF LIVERPOOL) 

(With Plates y-g) 

INTRODUCTION 

On the 27th of August 1949 the Fleetwood trawler Wyre General landed an unusual 
fish from the Bear Isle grounds. No information is available concerning the depth at 
which it was taken, but about 100 fathoms may be assumed from our knowledge of the 
fishery. Messrs. James Mitchell (Port Health Officer) and P. J. Fisher (Chief Sanitary 
Inspector), who have frequently been instrumental in obtaining rare fishes, kindly 
forwarded it to the Department of Zoology, University of Liverpool, where it was 
recognized as a rare Notacanthus and presented to the British Museum. The species is 
N. phasganorus Goode, new to the national collections. Only five other authenticated 
specimens are known, all in American museums, and of these but two have been des- 
cribed and figured. 1 

The holotype (U.S. National Museum, Washington, No. 25972 ; Goode (1881) ; 
Goode & Bean (1894 = 1896)) was taken from the stomach of a Ground-shark, Som- 
niosus brevipinna Lesueur = S. microcephalus (Bloch & Schneider), on the Grand 
Bank of Newfoundland, and was partly digested and mutilated about the head. 
Bigelow & Schroeder (1935) describe a specimen trawled in about 100 fathoms, 
20 miles south of Sable Island, which was in good condition except that the viscera 
had been removed, and the same authors mention a further example from the same 
locality (Museum of Comparative Zoology, Cambridge, Mass., Nos. 33946 and 35306 
respectively) . 

1 A large and originally well-preserved Notacanthus obtained in Iceland during the voyage of La 
Recherche and figured as N. nasus Bloch by Gaimard (1851, pi. XI) and by Cuvier (1836, pi. 55) has 
been tentatively referred to N. phasganorus Goode by Vaillant (18886), who was able to examine the 
specimen (Musee National d'Histoire Naturelle, Paris, No. A. 6864). One of us (D.W.T.) visiting Pans 
in October 1950 was told by Prof. L. Bertin that it could not then be found. 'Tres probablement a-t-il 
et6 detruit a une date ancienne (vers 1889) '. We have little doubt concerning the accuracy of Vaillant's 
identification, but do not regard the published figures and data available as sufficiently reliable for a 
critical determination. See Saemundsson (1949) for further discussion and a bibliography of Icelandic 
material. 



7 o ON NOTACANTHUS PHASGANORUS GOODE 

In reply to a request for further information on his material Dr. William C. 
Schroeder disclosed that two more examples have since been taken: M.C.Z. No. 37027 
in 420 fathoms at 42 18' N., 65 01' W., and No. 37037 in 100 fathoms at 44 N., 
57 W. Dr. Schroeder is preparing a paper on the species in which these will be de- 
scribed and has kindly allowed us to use such unpublished data as are needed to 
establish the identity of the Bear Island specimen. We wish also to acknowledge the 
assistance of Mr. Ernest A. Lachner of the U.S. National Museum who re-examined 
the holotype for us. The illustrations to the present paper are (with the exception 
of Fig. 1) the work of Mr. Hubert Williams and the X-ray photographs were taken by 
Mr. P. E. Purves. 

Modern papers by Matsubara (1938) on his Notacanthus fascidens and by Trotti 
(1939) on N. bonapartei Risso (based on the examination of 9 and 69 specimens 
respectively) have largely invalidated the taxonomic distinctions made by earlier 
workers, especially by Goode & Bean. Matsubara concludes: 

' Among the characteristics used in the taxonomy of the fishes of the family Notacanthidae, the 
number of anal spines and the positions of the insertions and also end points of the fins, which 
are in reality most variable, are considered to be of most importance. . . .It would be super- 
fluous to say that one must re-examine whether or not each known species belonging to the 
Notacanthidae is an independent species by taking the above mentioned variabilities into con- 
sideration.' 

Trotti remarks similarly : 

' Concludendo, la grande variability del profilo del muso e soprattutto la mancanza di persi- 
stenza del rapporto tra dorsali ed anali dure . . . ci porta ad una revisionedei caratteri differenziali 
dei rappresentanti del genere Notacanthus e Gigliolia.' 

In publishing this full account of the new specimen (British Museum (Natural 
History), No. 1950.3.30.2) we hope to put on record material of value to such a sub- 
sequent revision, and to justify an identification which not only extends the known 
range of N. phasganorus from the western Atlantic to the Arctic but also provides the 
first published data on the bionomics of the species if not of the genus. But although 
we now identify our specimen with Goode's species, we are conscious that in the 
present state of the taxonomy of the genus this name may not be final. There is need 
of a critical re-examination especially of the material designated N. chemnitzii Bloch 
1787, N. nasus Bloch 1795, N. phasganorus Goode 1881, and N. analis Gill 1883, the 
inter-specific differences between which, as at present described, do not seem greater 
than the infra-specific variation demonstrated elsewhere by Matsubara and by Trotti. 
It is probable that such a re-examination of the types of these four 'species' supple- 
mented by observations from other material will confirm our suspicion that some or 
all may be identical. This is no new speculation (see, for example, Liitken, 1898), and 
it may reasonably be inquired why no precise solution has yet been given. The answer 
is that apart from the comparative paucity of material, aggravated by its wide dis- 
persal in study-collections, even the type-locality of Bloch's material is not certainly 
established — though stated by him to have come from the East Indies it has since 
been believed to have come from Iceland — and the originally bad condition of the 
holotype has since further deteriorated. (Cf. accounts of Bloch himself, of Cuvier & 



ON NOTACANTHUS PHASGANORUS GOODE 



71 



Valenciennes (1831), and of Hilgendorf in Goode & Bean (1896).) Even if the specimen 
in the Berlin Museum is still in existence, it is therefore exceedingly doubtful whether 
it retains characters adequate for a modern redescription of Bloch's species. 

We have no more material relevant to that problem in the British Museum (Natural 
History), but hope in a subsequent paper to redescribe the types of N. sexspinis 
Richardson 1844 and N. annectens Boulenger 1904, and to give accounts of the series 
of these and related species in our collections as a contribution towards a future full 
revision. A forthcoming report on the Notacanthidae collected by the Danish Thor 
Expeditions in the north-eastern Atlantic will provide further material. 



DESCRIPTION 

Although the body is very well preserved, three factors seriously complicate the 
usual table of measurements. Firstly the fish is a spawning female, greatly distended 
by a mass of ripe eggs : as a consequence the vent is widely dilated, blocked by a large 
plug of ova, and opens posteriorly rather than ventrally, while the postero-lateral 
walls of the abdomen project as a pair of pouches which partly embrace the vent and 
conceal the origin of the spinous anal fin. This general distortion of the abdomen 
renders measurements of body-height of doubtful value. Secondly, the head of the 
specimen is markedly downturned in a very ' Mormyrid ' fashion and more so than in 
any figure or specimen of a Notacanthid that we have seen. Though there is little 
support for our opinion forthcoming from other specimens of N. phasganorus we are 
satisfied that the X-ray photograph published as Plate 8 and other considerations 
(dentition + diet, position of operculum in relation to gill-opening) indicate that this 
may at least be adopted as a natural attitude, even though it may not be the attitude 
of rest. Accordingly we give two measurements for body-length and other distances 
from the tip of the snout to various points; the first represents the measurements 
with the head forced into line with the body, the second with it in situ. Statements of 
body proportions are based on the former to facilitate comparison with other accounts ; 
the corresponding duplicate set may be computed from the data given if desired. 
Thirdly, there is some doubt regarding the tail, which may have had the tip broken 
off and subsequently regenerated a caudal fin. In this case it would be necessary to 
allow about another 5 cm. on the standard length, plus 2-3 cm. for the caudal fin. 



Measurements 



Total length 
Standard length 



970 mm. (950) 
945 „ (925) 



Body: 

Depth at pectoral 
,, pelvics 

vent 
Greatest depth 
Greatest breadth 
Length, snout to vent 



140 
170 
140 
180 

50 
422 



(402) 



72 



ON NOTACANTHUS PHASGANORUS GOODE 

Measurements (contd.) 



Head: 
















Length .......... 122 


mm 




Greatest depth ..... 








92 


,, 




Greatest breadth ..... 








5° 


,, 




Interocular .width ..... 








25 


,, 




Length of snout ..... 








35 


,, 




,, postorbital region . 








80 


,, 




,, upper jaw .... 








36 


,, 




mandible, to hind end of articular 








39 


,, 




Breadth of gape ..... 








4i 


,, 




Length of maxillary spine 








6 


,, 




Diameter of eye ..... 








21 


,, 




Longest gill-raker ..... 








6 


» 




Dorsal : 






Distance from snout ........ 352 


,, 


(35o) 


Length of base ......... 235 


,, 




Horizontal distance from pelvics . . . . . .12 


,, 




I II III IV V VI 


VII 


VIII IX X XI 


Lengths of spines . . . . .167889 


11 


12 10 13 14 mm 


Intervals between spines . . . 6 1520212423 




22 21 14 11 mm. 


Length of soft ray ......... 7 mm 




Anal: 






Distance from snout . -. . . . . . -432 


,, 


(412) 


vent. 










10 


,, 




Length of base .... 










540 


,, 




spinous base . 










230 


,, 




,, first spine 










2 


„ 




longest spine (XVIII) 










19 


,, 




,, ,, soft ray 










34 


» 




Pectoral : 






Distance from snout . . . . . . . .148 


)t 


(139) 


Length, left .......... 65 


,, 




right 56 


» 




Pelvic : 






Distance from snout ........ 350 


,, 


(33o) 


base to vent ....... 70 


,, 




"tip 2 4 


,, 




Length .......... 46 


» 




Caudal : 






Distance from tip to dorsal ....... 390 


„ 




Length ..... 










25 


,, 





Radial formula D. XI-i ; A. XX, 101+ ; C. I2( ?) ; P. 13; V. Ill, 7. 

Gill-rakers on first arch 3-f- 1 + 13. 

Branchiostegal rays 9. 

Vertebrae 185. (Nos. 75 and 80 have double centre.) 

(All counts from X-ray photographs.) 
Scales along lateral line, about 500. 

Scales in transverse series, 31 above lateral line, 58 below. 
Pyloric caeca destroyed through decomposition. 

Length of the head 7-95 times in the total length ; depth at pectoral 6-92 ; depth at 
pelvic 5 70; distance from tip of snout to dorsal 275; from tip of snout to pectoral 6-55; 



ON NOTACANTHUS PHASGANORUS GOODE 73 

from tip of snout to pelvic 277 ; from tip of snout to vent 2-29 ; tip of snout to anal 
2-24 ; from tip of caudal to dorsal 2-48 ; base of dorsal 4-12 ; spinous base of anal 4-21. 

Snout 3-48 in head ; eye 5-80 ; postorbital part of head 1-52 ; upper jaw 3-38 ; inter- 
ocular space 4-88 ; mandible 3-12 ; pectoral 1-87; pelvic 2-65. 

Body elongate, compressed, considerably higher at the pelvics than at the pectorals, 
even allowing for the distension of the abdomen ; the greatest breadth 0-35 the height 
at the vent ; tapering posteriorly into a long slender tail. 

Head compressed, shorter than depth of body, 2-46 in the trunk and 3-54 in the 
length from tip of snout to anal. Snout long, fleshy, 1-4 times the interocular width 
and i-66 times the diameter of the eye. Interocular space narrow, strongly convex, 
1-19 times the diameter of the eye. Eye covered by semi-transparent skin, lacking 
an orbital fold. Nostrils close together, much nearer eye than tip of snout, the 
posterior slit -like, one-third the eye's diameter from the orbit, the anterior opening 
into a thin-walled tube protected by a small flap. The centres of the eye, of the two 
nostrils, and the tip of the snout lie on a straight line. 

Mouth inferior, broad, gently curved ; upper jaw nearly as long as length of snout ; 
maxilla with a posteriorly directed pungent spine on its upper margin, extending to 
below the middle of the eye. The integument of the mandible forms a labial fold on 
each side. 

Teeth (PI. 7, fig. 4) in the upper jaw in a single row, 37 on each side, slender, 
inclining inward, the bases cylindrical, the tips antero-posteriorly flattened and in- 
trorse, mesially 3 mm. long, gradating into smaller and simpler lateral ones. Pala- 
tines movable vertically with two rows of about 25 rather finer teeth on each side, 
with sharper markedly introrse tips. Mandible with a complete innermost row of 
about 30 teeth on each side, resembling those of the upper jaw but more delicate, 
preceded by two irregular rows of fine aciculate teeth which do not extend as far 
laterally as those of the main series. All teeth more or less movable. Anteriorly the 
teeth of the upper jaw bite between the two series of the lower, but owing to the 
greater radius of curvature the posterior teeth bite outside those of the mandible. 
The palatine teeth engage with those of the lower jaw. No vomerine teeth. 

Gill-openings wide, membranes separate and free from isthmus. Gills four ; no 
pseudobranch visible on superficial examination. Gill-rakers slender, pointed, in- 
curved, well separated, having minute bristles on their inner faces ; a little more than 
half the length of the gill-filaments, the longest 3-50 in the diameter of the eye. 

The prominent pores of the lateralis system of the head are distributed thus : 3 in 
the supra-temporal series, and on each side 5 in the supra-orbital (comprising 2 
above the eye, 1 above the posterior nostril, 2 before the anterior nostril), 16 in the 
infra-orbital and 14 in the preoperculo-mandibular series. 

Lateral line gently arched over pectoral, following profile of the back, thence 
dropping obliquely to one-third the depth of the body over the vent, and further 
descending nearly to a median position at the point where it disappears two-thirds 
of the way along the tail. Lateral line pores conspicuous with darkly pigmented lips. 

Entire body scaled, even to the lips, except for the hinder margin of the opercu- 
lum. Scales cycloid, rectangulo-ovate, closely inset in tough sheaths; very small 
on the head (1-2 xro to 2-2 x 2-0 mm.), increasing in size posteriorly to a maximum 



74 ON NOTACANTHUS PHASGANORUS GOODE 

of 4*5x37 mm. on the middle of the body, and thereafter becoming progressively 
reduced until half-way along the tail they equal those of the head. 

Pectorals vertically inserted at middle of body-depth, at a distance behind the gill- 
opening equal to length of own base ; bases broad, fleshy, scaled, pedunculate ; pos- 
terior edge of fin rounded, length slightly more than half length of head. 

Pelvics (PL 7, fig. 3) closely adjacent, separated by a narrow groove, reaching far 
short of the vent. Bases fleshy, pedunculate, thickly covered with scales, origin very 
slightly behind vertical through origin of dorsal, posterior edge rounded. The third 
pelvic spine has two much smaller ones set against its base, the first of these concealed 
by skin. 

First dorsal spine (PL 9, fig. 6) hidden under the skin ; last dorsal spine the longest, 
followed by a recurved soft ray (PL 9, fig. 7) set in a fleshy protuberance. There is a 
slight groove between the bases of the spines and each supports a slight membrane 
posteriorly which is best developed between the last spine and the soft ray. 

The anal commences immediately behind the vent and below the Vth dorsal spine ; 
the XHIth anal spine lies under the last dorsal. The anal spines are embedded in 
fleshy tissue (the first completely concealed, PL 9, fig. 8), from which successive 
spines emerge farther and farther. 

Caudal (PL 9, fig. 9) clearly separated from anal, but lacking a distinct peduncle 
and probably aberrant owing to regeneration of tip (see p. 75). 

Colour. Head and body dark brown, tending to be fighter on the forehead and 
flanks ; lips and hinder edge of operculum bluish-black, fin-rays and anal fin dusky. 
The fish had a glossy, varnished appearance when dry. Peritoneum and stomach and 
inside of buccal cavity and operculum black, intestine cream. 

COMPARISON WITH SPECIMENS PREVIOUSLY DESCRIBED 

The original description of the holotype (Goode, 1881) gives the radial formula 
D. X ; A. XIX (130) ; Co ; P. (17) ; V. II, 8-9. Mr. Lachner was asked to re-examine 
the dorsal, pectoral, and spinous anal fins only, ascertaining whether any concealed 
spines and rays had been overlooked and whether a count of the pectoral rays obtained 
by means of an incision across the fleshy base required any modification of the above 
formula. He finds the right pectoral fin wanting and gives the count for the left : the 
revised formula now reads : 

Holotype: D. X-i ; A. XIX, 130; Co; P. 18; V. II, 8-9. 

compared with: 

M.C.Z. No. 33946 D. XI-(?); A. XXIV, 127; C. 7; P. 17; V. Ill, 7. 

New specimen, D. XI-i ; A. XX, 101+ ; C. i2( ?) ; P. 13 ; V. Ill, 7. 

Bigelow & Schroeder give A. XX for M.C.Z. No. 35306. Schroeder, in lit., provides 
the following additional data : 

M.C.Z. No. 35306 P. 16. One soft ray in dorsal. 
37027 P. 13. One 

37°37 P- J 6. Two soft rays in dorsal. 
>> 33946 Not available for re-examination. 

Bearing in mind the known variation in other species we may regard the counts for 



ON NOTACANTHUS PHASGANORUS GOODE 75 

dorsal, ventral, and spinous anal fins as giving an adequate agreement. 1 The range 
of variation in the pectoral (13-18) is remarkable, however, even compared with 
Trotti's counts for N. bonapartei (12-14) and Matsubara's for N. fascidens (12-15). 
The discrepancies in the counts given for the caudal in part reflect the curious mis- 
understanding which has surrounded the problem of the tail'structure in this group. 
The diagnoses of Goode & Bean (1894) contain mutual contradictions : 

Fam. N otacanthidae . 'Anal fin . . . extending ... to the caudal with which it unites.' 

Notacanthus. 'No caudal', although under the same generic diagnosis N. sexspinis is 
given a count of C. 5. In the accounts of the various species several numbers are given, 
including N. phasganorus with Co. 

Regan (1929) gives: 

Order Heteromi. ' A long tail, with a long anal fin below it, tapering to a point, without caudal 
fin.' 

While the relations of anal and caudal are certainly difficult to ascertain in these 
fishes and really call for radiographs and alizarin preparations for their proper eluci- 
dation, there can be no doubt that many previous descriptions made before the use of 
the binocular microscope became de rigueur will prove to be erroneous when the 
material is re-examined. 

The present specimen shows a distinct separation between the caudal and anal rays, 
more easily studied in an X-ray photograph (PL 9, fig. 9), which shows at least 
12 caudal rays. But the structure is markedly different from that of the tails of other 
species which we have examined, which are symmetrical, having a distinct though 
small caudal peduncle, already described and figured in N. phasganorus by Bigelow & 
Schroeder (1935). The appearance presented in our figure suggests that the tail has 
lost its tip at some time and subsequently regenerated a caudal fin. 

Since Goode almost certainly included the caudal rays in his count for the anal fin 
(130) we should do likewise to obtain a comparison, and so have 134 for the fish des- 
cribed by Bigelow & Schroeder and 113+ for the new specimen. A truncation of the 
tail would also account for this lower number. 

Gaimard's (185 1) figure of the La Recherche specimen evidently represents a tail 
even more markedly truncated (Vaillant, 1888b) and again with a regenerated caudal 
fin. It seems that this condition is not uncommon in Notacanthus. 

1 Vaillant's (1888&) data, supplemented by counts from Gaimard's (1851) plate, give the radial 
formula : 

D. XI-i; A. XXII, 92+; C. 8 (?); P. 16; V. Ill, 8 

for the La Recherche specimen, which therefore comes within tie known range of N. phasganorus. 

For further comparison the following counts all purport to have been taken on the holotype of N. nasus 
by Bloch (1795), Cuvier & Valenciennes (1831), and Hilgendorf for Goode & Bean (1896) respectively: 

D. X; A. + C. XIII, 136; P. 16; V. II, 8. 
D. X-O; A. XIII, 116; C. 8; P. 17; V. I, 8. 
D. XI; A. XV, 118; C. ?; P. 19; V. Ill, 7 (1), 8 (r). 

There seems to be little useful purpose in attempting to decide the relation between N. nasus and N. 
phasganorus on such data, except to remark that the only serious discrepancy, the consistently low count 
for the spinous anal, must be considered against the range of A. IX-XVIII demonstrated by Trotti 
(1939) in N. bonapartei, and the anterior fin-structure shown in our PI. 9, fig. 8. 

ZOOL. I. 5 K 



7 6 



ON NOTACANTHUS PHASGANORUS GOODE 
ANATOMY 



Those skeletal features discernible from X-ray photographs agree with the very 
full accounts given by Gunther (1887) for N. sexspinis and Vaillant (1888a:, b) for N. 
mediterraneus. Vaillant gives the more detailed account of the general anatomy. The 
viscera in the present specimen are in general poorly preserved, but it is possible to 
supplement these descriptions in certain details. 

The spacious body-cavity is very high, and extends posteriorly considerably 
behind the anus, to the level of the seventh anal spine. The kidneys are large, the 
deep anterior lobes flanking the rectum and not extending farther forward in any 



P. D.— , 




R.M. 



Fig. 1. Gas-bladder from left side. P.D., pneumatic duct ; A. and V., artery and vein supplying 
bladder; R.M., retia mirabilia. The dotted portions indicate the extensions of the pneumatic 
duct and of one rete within the bladder. 

bulk, while the remainder of the kidneys extend back along the roof of the post-anal 
body-cavity. There is no urinary bladder preserved. The undivided liver, the gonads, 
and the alimentary canal appear to agree with previous accounts, but the gas-bladder 
shows some marked differences and merits fuller treatment. Whether the dis- 
crepancies are due to interspecific variation or to inaccuracies of description cannot 
be stated. 

The gas-bladder (Fig. 1) is oval in shape with a small blind posterior prolongation, 
and lies above and extends slightly before the ventral fins. It is suspended in a fold of 
mesentery with a rather stronger attachment posteriorly ; the bulk of it being free 
anteriorly sags down into the body-cavity. The tunica externa comprises the usual 
two easily separable layers : an outer thin, tough, white, and muscular and an inner 
very dense and silvery, containing elastic fibres. The tunica interna comprises a sub- 
stantial basis of dense connective tissue supporting a poorly preserved series of 
muscular, vascular, and columnar epithelial layers. The lumen of the bladder con- 
tains a quantity of granular yellow matter. 

There is a fairly long pneumatic duct which does not approach anywhere near the 
oesophagus. Along it run the artery and vein supplying the bladder, and a number of 
streaks of yellowish tissue interpreted as pancreas. The vessels break up into two 
retia mirabilia before approaching the bladder with the pneumatic duct on the lower 



ON NOTACANTHUS PHASGANORUS GOODE 77 

left side, the combination of these structures forming a laterally compressed body 
which Gunther regarded as a left 'cornu' of the bladder, the retia evidently being 
identical with his 'pair of thick muscle-like pads'. The pneumatic duct opens in the 
centre of the floor of the bladder towards the anterior end. The retia are of the ' rete 
mirabile unipolare duplex' type (Woodland, 1911, 19110:), since dissection does not 
reveal any recombination of capillaries to form major vessels before they enter the gas- 
gland. The gas-gland is a small patch of spongy vascular tissue surrounding the en- 
trance of the pneumatic duct from which similar tracts radiate over most of the lining 
of the bladder. The postero-dorsal portion of the bladder has a thinner, smoother 
lining epithelium which probably represents a fully dilated oval (Woodland, 1913). 

BREEDING 

Though the precise date of capture is not available it may be assumed that the 
fish was taken about mid-August, and that the breeding season in Bear Island waters 
is therefore about that time. 

The ova, entangled in fibrous tissue, were opaque white when received and slightly 
elliptical, ranging from 1-20 X 1-30 down to 1-16 X 1-25 mm. diameter. They thus pro- 
vide a further instance of aspherical teleost eggs to be added to those discussed by 
Breder (1943). They contain many small colourless oil droplets, lo-jOfj, in diameter. 

FOOD AND FEEDING 

The stomach was well filled with the remains of some two dozen pink and magenta- 
coloured Actiniarians, comprising the tops of several small anemones of 1-2 cm. dia- 
meter and pieces apparently bitten from the rims of much larger ones. In some cases 
it was possible to distinguish scapus and scapulus ; all the fragments were more or less 
heavily tuberculated and bore traces of a dehiscent cuticle. 

A consideration of structure in relation to diet leads to some interesting conclusions. 

1. The dentition and shearing bite of the jaws are admirably suited to feeding on 
Actiniarians. What would, on theoretical considerations, seem the ideal shape of the 
head and position of the mouth ? A terminal mouth would require the fish to stand on 
end in the water when feeding, a rather unlikely proceeding, or to perform move- 
ments like those of the Lemon Dab Pleuronectes microcephalus Donovan which re- 
moves tubicolous polychaetes from their burrows by 'bringing its mouth down 
almost vertically upon its victim by a strong arching of the anterior part of the 
body' (Steven, 1930). (The same species in the southern North Sea feeds largely on 
Cerianthus sp. ; Todd, 1907.) This last movement is hardly possible to a stout- 
bodied fish such as our Notacanthus. There remains only the combination of an in- 
ferior mouth with what degree of flexure can be attained, the condition in fact which 
is illustrated in PL 8, where there is a marked downturning of the vertebral column 
bringing the jaws into the best position for horizontal and near-horizontal biting. 
From these considerations, accompanied by the fact that there is no indication of any 
fracture or dislocation of the skull and pectoral region, we believe that the head of our 
specimen is in fact being carried in a normal position, though whether this is faculta- 
tive or permanent cannot be decided. 



78 ON NOTACANTHUS PHASGANORUS GOODE 

2. The pieces of anemones present fall, as we have noted, into two size-groups, 
those from very small and very large individuals. The absence of remains of medium- 
sized ones suggests that such animals are possibly too large to be taken entire and yet 
too small to allow the fish to take a bite because the curvature of their body surface is 
so sharp that the jaws at maximum gape cannot obtain sufficient hold. With larger 
anemones it becomes possible to take a bite from the rim. 

3. Giinther (1887) remarks of N. sexspinis: 

' The osseous framework of this fish is so much wanting in the characteristic peculiarities of 
bathybial fishes as to throw serious doubts that this species at least of Notacanthus lives at a 
great depth.' 

The evidence from radiographs indicates that the skeleton of N. phasganorus is sub- 
stantially similar, and its gas-bladder is better developed than in oceanic fishes. But 
from its diet and the related structural adaptations it is clearly a bottom-feeding 
form, and it is therefore probable that specimens taken have been obtained on or near 
the bottom, so that a bathymetric distribution of 100 to at least 420 fathoms may be 
deduced from the records so far available. N. mediterraneus Fil. & Ver. is evidently 
another bottom-feeding form; Vaillant (18886) records hexactinellid sponge spicules 
from a specimen taken by the Talisman from more than 1,200 metres. 

Actiniarians have been reported as of frequent occurrence in Cod stomachs 
obtained from Bear Island and the Murman coast (Brown & Cheng, 1946) ; off 
Greenland, where Cod from deep water off Nuk feed almost entirely upon them 
(Jensen & Hansen, 1931), and in Danish waters (Blegvad, 1916). Stephenson, in 
Brown & Cheng, loc. cit., provisionally identified their material as Hormathia 
digitata (O. F. Mull.), H. nodosa (Fabr.), and Tealia felina (L.) var. lofotensis (Dan.). 
Some of our material may be referable to Hormathia spp., but precise identification 
would be extremely difficult if indeed possible. 

PARASITES 

The gills, alimentary canal, and peritoneum lining the body-cavity have been 
examined for parasites, but none have been found. 

REFERENCES 

Bigelow, H. B., & Schroeder, W. C. 1935. Two Rare Fishes, Notacanthus phasganorus Goode 
and Lycichthys latifrons (Steenstrup and Hallgrimsson) , from the Nova Scotian Banks. 
Proc. Boston Soc. Nat. Hist. 41: 13-18. 

Blegvad, G. 191 6. On the Food of Fish in the Danish Waters within the Skaw. Rep. Danish 
Biol. Sta. 24: 19-71. 

Bloch, M. E. 1787. Uber zwei merkwurdige Fischarten (Notacanthus chemnitzii und Silurus 
militaris). Abh. Bohm. Ges. Wiss. 3: 278-282. 

1795- Naturgeschichte der ausldndischen Fische, 9, Atlas: pi. 431. Berlin. 

Breder, C. M. 1943. The Eggs of Bathygobius soporator (Cuvier and Valenciennes) with a dis- 
cussion of other non-spherical Teleost Eggs. Bull. Bingham Oceanogr. Coll. 8 (3) : 1-49. 

Brown, W. W., & Cheng, C. 1946. Investigations into the Food of the Cod (Gadus callarias L.) 
off Bear Island, and of the Cod and Haddock (G. aeglefinus L.) off Iceland and the Murman 
Coast. Hull Bull. Mar. Ecol., 3, No. 18: 35-71. 



ON NOTACANTHUS PHASGANORUS GOODE 79 

Cuvier, G., & Valenciennes, A. 1831. Histoire Naturelle des Poissons, 8. Paris. 

Cuvier, G. 1836. Le Regne Animal ['Disciples' Edition']. Poissons. Paris. 

Gaimard, P. 1 85 1. Voyage en Islande et au Groenland. Zoologie, &c, atlas. Paris. 

Gill, T. N. 1883. Diagnosis of new genera and species of deep-sea fish-like vertebrates. Proc. 

U.S. Nat. Mus. 6: 253-260. 
Goode, G. B. 1 88 1. Notacanthus phasganorus. A new species of Notacanthide from the Grand 

Banks of Newfoundland. Proc. U.S. Nat. Mus. 3: 535-537. 
& Bean, T. H. 1894. A Revision of the Order Heteromi, deep-sea fishes, with a description 

of the new generic types Macdonaldia and Lipogenys. (Sci. Res. Albatross Exped. XXIX.) 

Proc. U.S. Nat. Mus. 17: 455-470. 

1896. Oceanic Ichthyology. Mem. Haw. Mus. Comp. Zool. 22. 

Gunther, A. 1887. Report on the Deep-Sea Fishes. Rep. Sci. Res. 'Challenger', Zoology, 22. 
Jensen, A. S., & Hansen, P. M. 193 1. Investigation on the Greenland Cod (Gadus callarias L.). 

Rapp. Cons. Explor. Mer. 72. 
Lutken, C. 1899. Danish Ingolf Exped. 2, Pt. I. 1-39. Copenhagen. 
Matsubara, K. 1938. Studies on the Deep-Sea Fishes of Japan. X. On a New Fish, Notacanthus 

fascidens, belonging to Heteromi with Special Reference to its Variations. Bull. Jap. Soc. 

Sci. Fish. 7: 1 31-136. 
Regan, C. T. 1929. Fishes. [Article in] Encyclopaedia Britannica, 14th edn: 317. 
Saemundsson, B. 1949. Marine Pisces. Zoology of Iceland. Ed. A. Fridriksson & S. L. Tuxen. 

4 (72). 
Steven, G. A. 1930. Bottom Fauna and the Food of Fishes. /. Mar. Biol. Ass. U.K. n.s. 16: 

677-700. 
Todd, R. A. 1907. Second Report on the Food of Fishes (North Sea, 1 904-1 905). Rep. (Southern 

Area) Fish Invest. N. Sea, 1904-1905 (Mar. Biol. Ass.), 2 (1) : 48-163. [Ed. 3837.] 
Trotti, L. 1939. Contributo alia Conoscenza del Genere Notacanthus ed in particolare della 

Specie Bonapartei Risso. Ann. Mus. Stor. Nat. Genova, 60: 363-378. 
Vaillant, L. 1888a. Sur les rapports zoologiques du genre Notacanthus Bloch. C. R. Acad. Sci. 

107: 75 1 -753- Paris. 

1888&. Exped. Sci. 'Travailleur' et 'Talisman'. Poissons. Paris. 

Woodland, W. N. F. 191 i. On the Structure and Function of the Gas Glands and Retia 

Mirabilia associated with the Gas Bladder of some Teleostean Fishes, with Notes on the 

Teleost Pancreas. Proc. Zool. Soc. Lond. 1911: 183-238. 
— — 1911a. On some Experimental Tests of Recent Views concerning the Physiology of Gas 

Production in Teleostean Fishes. Anat. Anz. 40: 225-242. 
1 91 3. Notes on the Structure and Mode of Action of the 'Oval' in the Pollack (Gadus 

pollachius) and Mullet (Mugil chelo). J. Mar. Biol. Ass. U.K. 9: 561-565. 



V 



PRESENTED 

1- N0< 



PLATE 7 

Fig. 2. Notacanthus phasganorus Goode; Bear Island specimen. 

Fig. 3. Detail of right pelvic fin, from below. 

Fig. 4. A, underside of head; B, side, and C, D, front views of teeth of maxillary series; E. 
palatine tooth. 



ull. B.M. [N.H.) Zoology, I. 5 



PLATE 7 






in 
D 

O 

< 

o 

< 

a p 
w 

H 

o 

H 
O 



PLATE 8 
Fig. 5. Unretouched X-ray photograph of head, showing flexure of vertebral column. 



Bull. B.M. {N.H.) Zoology, I. 5 



PLATE 8 




1 



Fig. 5 
NOTOCANTHUS PHASGANORUS 



PLATE 9 

Fig. 6. X-ray photograph of origin of dorsal fin. 

Fig. 7. X-ray photograph of end of dorsal fin. I, II, &c, spines ; R, soft ray. 

Fig. 8. X-ray photograph of pelvic region, showing pelvic fins and girdle. AI, first spine of 
anal fin. 

Fig. 9. X-ray photograph of end of tail. 

(Figs. 2-4, scale indicated on drawing ; Figs. 5-8, x 1 ; Fig. 9, X 2.) 



Bull. B.M. (N.H.) Zoology, I. 5 



PLATE 9 



w n 



n i 




Fig. 6 




Fig. 




Fig. 




Fig. 9 
NOTOCANTHUS PHASGANORUS 




PRESENTED 

NOV 1951 



PRINTED IN 

GREAT BRITAIN 

AT THE 

UNIVERSITY PRESS 

OXFORD 

BY 

CHARLES BATEY 

PRINTER 

TO THE 

UNIVERSITY 



J 




3 1 OCT 1951 

THE 

OCEAN SUNFISHES 
(FAMILY MOLIDAE) 



A. FRASER-BRUNNER 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ZOOLOGY Vol. i No. 6 

LONDON : 195 1 



THE OCEAN SUNFISHES 
(FAMILY MOLIDAE) 



BY 



A. FRASER-BRUNNER 



1 




Pp. 87-121 ; 18 Text-figures 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ZOOLOGY Vol. 1 No. 6 

LONDON : 1951 



THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted in 1949, is to be 
issued in five series, corresponding to the Departments 
of the Museum. 

Parts will appear at irregular intervals as they 
become ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

This paper is Vol. 1, No. 6 of the Zoology series. 



PRINTED BY ORDER OF THE TRUSTEES OF 
THE BRITISH MUSEUM 



Issued November 1951 Price Seven Shillings and Sixpence 



THE OCEAN SUNFISHES (FAMILY MOLIDAE) 

By A. FRASER-BRUNNER 

SYNOPSIS 

The relationships of the Molidae with other Plectognathi are briefly discussed. The movable lobe at the 
hind margin of the body, supported usually by migrant dorsal and anal rays but sometimes also by 
caudal rays centrally, is designated the ' clavus '. Three genera are recognized, assigned to two subfamilies. 
Masturus is shown to include two forms (treated as species but possibly the sexes of one). Evidence is 
presented to show that in this genus alone of the family some caudal rays are developed. Mola is shown 
to include two species, which are diagnosed and figured. Sexual dimorphism in Mola mola is described. 
Full synonymies are included. 

ON account of their curious form and the great size which they often attain, the fishes 
of the family Molidae, usually called Ocean Sunfishes, have attracted considerable 
attention from early times. A large and scattered literature exists concerning them, 
but although comparative studies have been made from time to time and their 
anatomy has received attention quite frequently, we are still far from a complete 
understanding of their relationships. This is mainly because all the species are rather 
rare, and their occurrence unpredictable, so that it is not possible to make an excur- 
sion for the express purpose of collecting specimens, as could be done with many other 
fishes, while the great size of most examples makes transportation and preservation 
a difficult problem. Consequently good comparative material is not easily available 
for study, and much reliance has to be placed upon published descriptions and figures. 

It is the purpose of the present work to draw attention to certain facts which have 
become apparent from a study of the literature, aided by the material in the national 
collection. 

My thanks are due to Mr. G. Palmer for his assistance in seeking out some of the 
references and checking a number of points in them. 

I am concerned here only with taxonomy within the family, since a full considera- 
tion of their relationship to other Plectognathous fishes will be included in a larger 
work upon the anatomy and phylogeny of the whole Order now in preparation. It 
can be pointed out here, however, that I have already indicated in an earlier paper 
(Fraser-Brunner, 1943), that the Molidae are not really as highly specialized as 
previously supposed. Their main peculiarity lies in the atrophy of the rear end of the 
vertebral column, resulting in a mechanical rearrangement of the median fin- 
structures closely resembling that seen in other fishes when the tail is amputated al 
an early age ; some interesting examples of this among Flatfishes have been given by 
Chabanaud (1935). The resemblance is not quite perfect, since with amputation the 
supporting bones of dorsal and anal fins are lost with the tail, whereas in the Molidae 
only the vertebral structures are lost. 

The lateralis muscles of the trunk, deprived of their normal attachment, become 
inserted upon the deep muscles of the dorsal and anal fins, and progressively lose 
their identity in the genera Ranzania, Masturus, Mola. The result of this is that body- 
flexion is lost but the dorsal and anal fins gain in power, and the latter are therefore 



9 o THE OCEAN SUNFISHES (FAMILY MOLIDAE) 

the principal means of locomotion. The posterior parts of these two fins extend round 
the rear end of the truncated body to support a broad, stiff lobe which acts as a 
rudder. This has been called the 'pseudo-caudal' by Raven (1939 a), but this is not 
a very suitable term in my opinion ; any structure in this part of the body may be 
described as 'caudal', and even if 'pseudo-caudal fin' is used, this is not true for all 
the species, for I hope to demonstrate on a later page that remains of the true caudal 
fin are included in the structure in Masturus. 

For this rudder-like lobe at the end of the body in the Molidae I therefore propose 
using a new term, and throughout this paper it will be called ' the clavus ' (Lat. clavus, 
a rudder) . 

Apart from these changes of form, all of which are demanded as mechanical 
consequences of the phyletic atrophy of the posterior part of the vertebral column, 
the Sunfishes resemble in their anatomy the more primitive of the Tetraodont fishes, 
and in one feature at least, the retention of the fourth gill, they are less modified even 
than those. They stand, therefore, near the main stem of the Tetraodonts, and 
attempts to derive them from the highly modified Diodontidae seem to me to be very 
far-fetched ; whatever resemblances the latter may show are more plausibly explained 
by the assumption that they are evolved from a Mola-like type (before caudal 
atrophy) rather than the reverse. The Molidae show also some features in common 
with the Ostraciontoidea alone among Plectognathi, and indicate therefore the diver- 
gence of the Trunkfishes and Puffer-fishes from a common stock during their evolu- 
tion. 

In my classification of the Tetraodontoidea I expressed the view that only two 
genera of Molidae should be recognized. This was based on the belief, current at that 
time, that Ranzania, Masturus, and Mola were each represented by a single species, 
and since the latter two forms seemed to be more closely related to each other than 
to Ranzania, it appeared that this relationship would be better expressed by placing 
them together in the genus Mola. A more detailed examination of these fishes, 
however, has caused me to modify these views. 

Firstly, I find that there are two species of Mola in the limited sense — one of world- 
wide distribution and the other apparently restricted to the Australasian region. 
Whitley (1931) recently revived the name Mola ramsayi Giglioli 1883 for the latter, 
but was apparently unaware of its distinguishing characters and assumed that all 
specimens from that region should be so named, whereas his main description appears 
to be of M. mola and the records show that both M. mola and M. ramsayi are to be 
found around Australasia. The type of M. ramsayi, a huge stuffed specimen, is in the 
British Museum (Natural History) , and by a piece of good fortune one of our spirit- 
specimens belongs to that species, so that I have been able to make direct comparison 
with examples of M. mola of similar size. 

Secondly, a close study of the literature concerning Masturus lanceolatus, aided 
considerably by the excellent work of Gudger on this subject, reveals that two forms 
are included here also, though it is not certain that they are different species. More 
interesting is the apparent fact that in Masturus alone of the family a remnant of the 
caudal fin is included in the support of the clavus. In this and in its musculature it is 
a little less specialized than Mola, and it therefore now seems desirable to recognize 



THE OCEAN SUNFISHES (FAMILY MOLIDAE) 91 

it as a distinct genus in order to express its relationship to the other genera more 
clearly. 

There has been much speculation in the past as to whether the rays supporting the 
clavus belong to the caudal fin or to the dorsal and anal, and even Gregory & Raven 
(1934), when describing the anatomy of M. mola, thought them to be caudal although 
their description and figure indicate that they are not (an error corrected by Raven 
in 1939). Apart from internal anatomy, the number of these rays is in most cases 
against the likelihood that they all belong to the caudal fin ; in most Plectognathi 




Fig. 1. Diagram illustrating reduction of the caudal region in the Molidae. 

Persistent parts of the axial skeleton shown in black ; atrophied parts shown with broken 

line; the last interneural and interhaemal bones close in along the lines marked with 

arrows. (Based on Ryder, 1886, modified by reference to adult and larval forms.) 

there are only 12 caudal rays, exceptionally 13, and sometimes as few as 10. But in 
Ranzania and Masturus the clavus is supported by 20 or more rays, in Mola ramsayi 
by 16, and only in Mola mola by 12. 

These rays are, in the main, supported by elements which have all the appearance 
of belonging to the series of interspinous supports of the dorsal and anal fins, but have 
been rotated to lie roughly at right angles to the last normal vertebro-interspinous 
complex by the process which has already been suggested by Ryder (Fig. 1) . The 
skeletal supports of the clavus are accompanied by muscles which have split off from 
the inclinators of the dorsal and anal fins, and caudal muscles appear to have been 
lost with the posterior vertebral structures. Reduction of the caudal region can be 
shown to extend to the number of rays supporting the clavus. Thus in Ranzania, 
which has 18 remaining vertebrae, there are 22 rays of dorsal and anal origin in the 
clavus ; in Masturus and Mola, which have 17 vertebrae, we find the series : Masturus 
14-18 (exclusive of caudal rays), Mola ramsayi 16, M. mola 12. 

Alongside this the form of the rays is of interest (see Barnard, 1935). In Ranzania 



9Hofa 



92 THE OCEAN SUNFISHES (FAMILY MOLIDAE) 

each ray in the clavus (except the outermost) is abruptly branched at its distal end 
(like those of the dorsal and anal lobes) and forms a fairly stiff fan-shaped support, 
closely apposed to those each side of it. In Mola this branched portion becomes 
hyperossified into a single plate or ossicle characteristic of the genus, the number and 
arrangement of these ossicles being of importance in specific diagnosis. 

In Masturus the rays seem never to be branched in adults, and are never ossified 
distally, but in young examples they may be branched at the tip like those of Ran- 
zania. This development can be seen by comparing the figures accompanying this 
paper. Between the rays in Ranzania lie elongate lobes of apparently collagenous 
material (shown in Fig. 3), and it is probably these which in Mola mola extend back 

between the widely spaced rays to form the 
lobes characteristic of the clavus of large speci- 
mens of that species. 

As a matter of interest, it may be remarked 

that Ranzania, Mola, and the Percomorphous 

family Carapidae (Fierasferidae) are the only 

fishes to which the term ' gephyrocercal ' can 

properly be applied, as pointed out by Ryder 

, \ / when originally proposing the term. 

/&CMWCCfttCt\ / Raven has taken the view that Ranzania is 

^ \ / the most specialized of these genera. I cannot 

agree with this. Its skeleton is much less de- 
generate than that of Mola, more strongly ossi- 
fied, and there are 18 or 19 vertebrae. The 
lateralis muscles are still moderately developed, 
though inserted posteriorly on the m. dorsalis 
profundus ; the usual division of the dorsal por- 
tion into superior and inferior parts is still quite 
distinct anteriorly. I feel sure that Raven was 
mistaken in identifying the lateralis muscles as 
dorsal and anal depressors ; they insert on to the 
latter but are distinct. The gill-rakers are free 
and apparently functional as in more generalized fishes. Further, this species is not 
gigantic. 

It is not suggested, however, that Ranzania is completely representative of the 
ancestor of the other two genera ; it has retained more primitive features, but it has 
completely lost the caudal fin, whereas Masturus, which is otherwise a stage farther 
towards Mola, retains a vestige of this fin, as will be shown later. 

The relationships of these genera are therefore probably as shown in Fig. 2. An 
ancestral form in which the skeleton and musculature is still fairly normal and the 
caudal fin not completely lost gives rise to Ranzania on the one hand, which loses its 
caudal fin, and to Masturus on the other, in which the caudal fin retains a precarious 
hold but the skeleton and the musculature deteriorate. Further degeneration and 
complete loss of the caudal fin in this second line gives us Mola. 

The need to recognize Masturus and Mola as more closely related to each other 




Fig. 2. Relationships of the genera of 
the family Molidae. 



THE OCEAN SUNFISHES (FAMILY MOLIDAE) 93 

than to Ranzania therefore still remains, and these two lines can now be expressed as 
subfamilies. 

The three genera illustrate quite well the manner in which the lateralis muscles lose 
their primary function of flexing the body and become successively more closely 
associated with the dorsal and anal fins, their added power enabling these fins to 




Fig. 3. 



Ranzania laevis, adult. A specimen 515 mm. long, from Baltimore, 
County Cork, Ireland. 



become proportionately larger. The body is therefore held rigid, assisted in 
Ranzania by a carapace similar to that of Strophiurichthys among the Ostracionts, 
but with much smaller hexagonal plates; in Mashirus and Mola this carapace is 
reduced to small denticles, and rigidity is assisted by a thick collagenous layer 
beneath the skin (Green, 1901). 

All the species pass through a remarkable metamorphosis. The newly hatched 
larvae are Tetraodon-hke, but soon (at 1-8 mm.) develop a cuirass of broad plates 
with jutting triangular projections, looking more reminiscent of an Ostraciont 
(Richardson named this stage Ostracion boops). With the atrophy of the larval tail, 
Ranzania seems to pass, by reduction of the cuirass and elongation of the body, into 



94 THE OCEAN SUNFISHES (FAMILY MOLIDAE) 

a form essentially like that of the adult though proportionately longer, but Masturus 
and Mola show an intermediate stage, wherein the cuirass breaks up into small 
denticles and the triangular projections grow into long sharp spines on broad poly- 
gonal grooved bases (very like those of Acanthostracion or Lactoria). This stage is 
much shorter in the body than the adult. As growth proceeds the body lengthens and 
the spines shorten and disappear, though in Mola the bases of one on the snout and 
one at the chin are nearly always retained even in the largest specimens. 

KEY TO THE GENERA OF MOLIDAE 

I. Form comparatively elongate. Vertebrae 8+10 or 11. Carapace of smooth hexa- 
gonal plates, 1 terminating at bases of dorsal and anal fins and clavus. Lips pro- 
duced forward beyond teeth as a funnel, closing as a vertical slit. Gill-rakers free. 
2 uppermost branchiostegal rays coalesced. Clavus with 22 rays, all borne 
on interspinous bones. No secondary post-larval metamorphosis (subfamily 
Ranzaniinae) . . . . . . . . 1. Ranzania 

II. Form shorter. Vertebrae 9+8. Carapace collagenous; skin of body and clavus 
with small rough denticles. Lips not funnel-like. Gill-rakers concealed in thick 
skin. 6 distinct branchiostegal rays. A secondary post-larval metamorphosis 
(subfamily Molinae). 

A. Median rays of clavus not borne on interspinous bones, supporting 
a pronounced lobe; none of the rays bearing ossifications 
distally . . . . . . . . .2. Masturus 

B. All rays in clavus borne on interspinous bones, most of them terminating 
in an ossification distally ...... 3. Mola 

Genus RANZANIA Nardo 

? Triurus Lacepede, 1800, Hist. Nat. Poiss. 2: 200. Type: Triurus bougainvillianus Lacepede. 

Ranzania Nardo, 1840, Ann. Sci. Regno Lombardo-Veneto, 5: 10, 105. Type: Ranzania typus 
Nardo (= Ostracion laevis Pennant). 

Centaurus Kaup, 1855, Arch. Naturgesch. 21 (1) : 221. Type: Ostracion hoops Richardson (= Os- 
tracion laevis Pennant, young). 

The characters of this genus have been indicated concisely in the foregoing key. 

Lacepede's description of Triurus bougainvillianus was based upon manuscript 
notes by Commerson. It could be interpreted as referring to the fish later known as 
Ranzania, but to describe the funnel-like lips as 'rictu fistulari' or ' le museau avance 
en forme de tube' and again 'un museau tres prolonge fait en forme de tube assez 
etroit' requires a good stretch of imagination. Moreover, the depth of the body is 
given as the proportion of 18 against the body-length of 71, and no other Sunfish has 
been recorded as slender as that. Finally, it has to be noted that in vol. 1 of the same 
work Ranzania is figured (pi. 22) under the name 'le Tetrodon lune\ The status of 
the name Triurus is therefore doubtful, and I hesitate to follow Whitley in using it, 
particularly since the name Ranzania is so well known. 

A single species. 

1 In young specimens (90 mm.) each plate has a prominent bony tubercle centrally. 



THE OCEAN SUNFISHES (FAMILY MOLIDAE) 95 

Ranzania laevis (Pennant) 

Ostracion laevis Pennant, 1776, Brit. Zool, ed. 4, 3: 129, pi. 19. 

Tetrodon truncatus Retzius, 1785, K. Svenska Vetensk Akad. Handl. 6 (2) : 121 (based on Pennant) ; 
Lacepede, 1798, Hist. Nat. Poiss. 1: 514, pi. 22 f. 2; Donovan, 1808, Nat. Hist. Brit. Fish. 
2: pi. 41. 

Orthragoriscus oblongus Bloch & Schneider, 1801, Syst. Ichth.: 511, pi. xcviii. 

Orthagoriscus oblongus Jenyns, 1835, Man. Brit. Vertebr. Anim.: 491 ; Yarrell, 1836, Hist. Brit. 
Fish. 2: 357, fig.; Couch, 1841, Ann. Mag. Nat. Hist. 6: 144; Bonaparte, 1846, Cat. Met. 
Pesci eur.\ 88; Bleeker, i860, Natuurk. Tijdschr. Ned.-Ind. 21: 57; Couch, 1865, Hist. Fish. 
Brit. Is. 4: 381, pi. 246 ; Harting, 1868, Verh. Akad. Wet. Amst. : 12, pi. 2, fig. 2 ; Andrews, 1871, 
Proc. Nat. Hist. Soc. Dublin, 5: 123; Sauvage, 1891, Hist. Madagascar, 16 (Poiss.): 529; 
Nobre, 1935, Faun. Mar. Portugal, Vertebr.: 242. 

Cephalus oblongus Shaw, 1806, Gen. Zool. 5: 439, pi. 176; Swainson, 1839, Nat. Hist. Class. Fish. 
2: 330. 

Cephalus varius Shaw, 1806, ibid. 

Orthragus commersoni Rafinesque, 1810, Caratt. Sicilia: 18. 

Orthragus oblongus Rafinesque, 1810, Indice Itt. Sicil.: 40. 

Tetraodon truncatus Couch, 1825, Trans. Linn. Soc. Lond. 14: 88. 

Cephalus elongatus Risso, 1826, Eur. Merid. 3: 173. 

Mola planci Nardo, 1828, Bull. Sci. Nat. Ferussac, 13: 437. 

Orthagoriscus truncatus Fleming, 1828, Hist. Brit. Anim.: 175; Gunther, 1870, Cat. Fish. Brit. 
Mus. 8: 319; Bleeker, 1873, Ned. Tijdschr. Dierk. 4: 121 ; 1879, Verh. Akad. Wed. Amst. 18: 26; 
Rochebrune, 1883-1885, Faune Senegambie (Poiss.): 157; Day, 1884, Fish. Gt. Brit.: 276, 
pi. 149; Beauregard, 1893, Bull. Soc. Sci. Nat. Ouest, 3: 229; Scharff, 1906, Irish Nat. 15: 275 ; 
Mauro, 1906, Boll. Accad. Gioenia, Catania, n.s. 85: 16, fig. 

Cephalus cocherani Traill, 1832, Mem. Werner: 6. 

Orthragoriscus elegans Ranzani, 1839, Novi Comment. Acad. Sci. Inst. Bonon. 3: table. 

Orthragoriscus battarae Ranzani, 1839, ibid. 

Ranzania typus Nardo, 1840, Ann. Sci. Regno Lombardo-V eneto , 5: 105; Smith, 1949, Sea Fish. 
S. Afr.: 422, pi. 95, fig. 1212. 

Ostracion boops Richardson, 1844, Voy. Erebus and Terror, Fish. : 52, pi. 30, figs. 18-21 ; Gunther, 
1880, Intro. Study Fish.: 175, fig. 93. 

Orthagoriscus planci Bonaparte, 1846, Cat. Met. pesci eur.: 88; Canestrini, 1872, Fauna d'ltalia, 
Pesci: 149; Stossich, 1879, Boll. Soc. Adriat. Sci. Nat. 5: 36. 

Orthragoriscus lunaris (Gronow) Gray, 1854, Cat. Fish.: 165. 

Centaurus boops Kaup, 1855, Arch. Naturgesch. 21 (1): 221. 

Ranzania truncata Jordan & Gilbert, 1883, Bull. U.S. Nat. Mus. 16: 966; Trois, 1884, Atti 1st. 
Veneto, 2 (6) pt. 1: 1269, pis. 12-14; pt- 2: 1543, pi. 16; Perugia, 1897, Ann. Mus. Stor. nat. 
Genova (2), 18: 140 ; Jordan & Evermann, 1898, Bull. U.S. Nat. Mus. : 47 (2) : 1755 ; Steenstrup 
& Liitken, 1898, K. danske vidensk. Selsk. Skr. (6) 9: 54, fig. ; Gunther, 1910, /. Mus. Godeffroy, 
9 (17) : 477; Pellegrin, 1912, Bull. Soc. Zool. France, 37: 228, fig. 1 ; Ribeiro, 1915, Arch. Mus. 
nac. Rio de J. 17 (Molidae) : 4, pi. ; Thompson, 1918, Mar. Biol. Rep. Cape Town 4: 176 ; Buen, 
1919, Bol. Pesc. Madr. 4: 295; 1935, Notas Inst. esp. Oceanogr. 2 (81): 146; Schmidt, 1921, 
Nature, Lond. 107: 76, figs. 2, 4, 5 ; Medd. Komm. Havundersog. Fisk. 6 (6), fig. 2. 13, pi. 1, 
fig. 7 ; Fowler, 1922, Copeia 112: 84 ; Vinciguerra, 1923, Comune di GenovaZ: 5, fig. 3 ; Barnard, 
1927, Ann. S. Afr. Mus. 21: 989, fig. 32; Fowler, 1928, Mem. Bishop Mus. 10: 475; Schmidt, 
1932, Dana's Togt omkr. Jord.: 251, fig. 197 (6-1 1); Gudger, 1935, Nature, Lond. 135: 548; 
Barnard, 1935, Ann. S. Afr. Mus. 30: 655, fig. 6c; Ehrenbaum, 1936, Handb. Seefisch. Nord- 
europ. 2: 88, fig. 69; Gudger, 1936, Nature, Lond. 137: 947; Fowler, 1936, Bull. Amer. Mus. 
Nat. Hist. 70 (2): 1123, fig. 470; Ninni, 1939, Atti. Soc. Hal. Sci. nat. 78: 236; Raven, 1939, 
Amer. Mus. Novit. 1038, figs. 1-3; Clark, 1949, ibid. 1397: 7, fig. 9; Maul, 1949, Vertebr. 
Madeira 2 (Peixes) : 158. 

Ranzania makua Jenkins, 1895, Proc. Calif. Acad. Sci. (2) 5: 780, pi. ; Fowler, 1900, Proc. Acad. 
ZOOL. I. 6 M 



9 6 



THE OCEAN SUNFISHES (FAMILY MOLIDAE) 



Nat. Sci. Philad.: 514; Jordan & Snyder, 1901, Proc. U.S. Nat. Mus. 24: 262; Jenkins, 1902, 

Bull. U.S. Fish. Comm. 22: 486 (1903) ; Jordan & Evermann, 1905, Bull. U.S. Fish. Comm. 

23 (1): 439, fig. 194; Jordan, Tanaka, & Snyder, 1913, /. Coll. Sci. Tokyo 33: 231, fig. 166; 

Snyder, 1913, Proc. U.S. Nat. Mus. 44: 460, pi. 63; Tanaka, 1914, Fig. Descr. Fish. Japan 

16: 274, pi. 76; Jordan & Jordan, 1922, Mem. Carneg. Mus. 10 (1) : 89; Jordan, Evermann, & 

Tanaka, 1927, Proc. Calif. Acad. Sci. 16 (4) : 680. 
Orthagoriscus (larva) Sanzo, 1919, Mem. R. Com. Talassogr. ital. 69: 1-7, figs. 1-4. 
Ranzania laevis Whitley, 1933, Vict. Nat. 49: 211, figs. 6, 7 ; Phillips, 1941, Trans. Proc. Roy. Soc. 

N.Z. 71 (3) : 245, pi. 41, fig. 6; Deraniyagala, 1944, /. Bombay Nat. Hist. Soc. 44 (3) : 429. 
Triurus laevis Whitley, 1937, Mem. Queensland Mus. 11 (2): 147; Hale, 1944, S. Aust. Nat. 22: 

pt. 4, pi. 1, figs. 





Fig. 4. Ranzania laevis. Front view of head, showing mouth open and closed. 



Examination of the records leaves little doubt that a single species of Ranzania 
ranges the seas of the whole world except the polar regions, but it seems that two 
subspecies can be recognized as follows : 

Ranzania laevis laevis (Pennant). Depth of carapace contained twice or more in 
its length, in adults (up to 580 mm.). Axil of pectoral fin well below level of 
centre of eye. Height of anal fin less than § length of head. Atlantic Ocean. 

R. 1. makua Jenkins. Depth of carapace contained less than twice in its length, 
in adults (400-500 mm.). Axil of pectoral fin above level of centre of eye. 
Height of anal fin f length of head or more. North Pacific Ocean. 

That these two forms are simply subspecific extremes in the range is shown by the 
records from the Indian Ocean, wherein the depth is usually given as for makua while 



THE OCEAN SUNFISHES (FAMILY MOLIDAE) 97 

the pectoral fin is low as in laevis. A specimen from Mauritius in our collection shows 
these features well, and a closely similar specimen has been figured by Whitley from 
Australia. 

Whenever the coloration has been described it has been shown to be closely similar 
in all these forms, a pattern of pale transverse bands on a darker ground, the bands 
edged with spots and broken lines of black ; three bands associated with the eye are 




Fig. 5. Development of Ranzania laevis. 

A, larva (1-7 mm.) ; B, C, D, early, full, and late ' Ostracion hoops' stage ; E, transition to adult form 

(8 mm.). (After Schmidt.) 



the most constant, the posterior ones being variously broken or anastomosed, some- 
times enclosing large oval areas of the ground colour. The colours are said to be very 
brilliant in life. 

The mouth is very curious, the lips extending well beyond the teeth and forming a 
funnel, the mouth being oval with the long axis vertical. The orifice closes along this 
axis, so that the rictus is really vertical — apparently unique among fishes (Fig. 4). 
This feature was shown clearly in the earliest published picture of the fish (Aldro- 
vandi, 1613), a remarkably good representation for its period. 

Too little is known of the feeding habits to show whether they can be associated 
with the peculiar mouth, but the species has been reported to take littoral algae 



98 THE OCEAN SUNFISHES (FAMILY MOLIDAE) 

(Barnard, 1927), and it is possible that the lips can suck in and close upon a frond 
while the teeth nip it off. 

The fine developmental series given by Schmidt (1932) shows that Raven was 
correct in supposing that elongation of the body is secondary, but it also shows 
that Ranzania is never so greatly shortened as the other two genera (Fig. 5). 
Lengthening occurs after the 8-mm. stage, until at 53 mm. the length of the carapace 
is about 3 times its depth. This proportion is maintained up to 90-mm. size, and after 
that the depth of the body increases with growth, so that at 250 mm. the length of 
the carapace is 2-5 times its depth, at 430 mm. 2-25 times, at 515 mm. 2-1 times, 
and at 580 mm. only twice. These figures are for the Atlantic form as shown by our 
specimens ; in the North Pacific subspecies R. makua either the early lengthening is 
not so great or the later deepening is more rapid. 

The general use of the name truncatus for this species seems to date from Gunther, 
1870 ; it is not clear why he chose this name rather than that of Pennant, on whose 
work that of Retzius was based, but possibly it was due to the fact that Pennant's 
description was numbered 54, while on his plate the number 54 appears beside a 
figure of the ' Short Diodon ' (Mola mola), leaving the other Sunfish without a number. 
As both description and figure are titled 'Oblong Diodon', however, this is clearly 
an error in numbering, and there can be no doubt as to the identity of Ostracion 
laevis, which antedates Tetrodon truncatus by nine years. 

Ranzania laevis does not reach so enormous a size as the other members of the 
family, apparently not exceeding 800 mm. in length. It has been recorded from all 
warm seas, as far north as Scandinavia and far south as New Zealand, usually from 
single specimens — though it was once observed in great numbers at the surface of 
the water off Martinique (Pellegrin, 1912). As Schmidt has pointed out, most records 
of larval Sunfishes to date belong to this species, and he has given us a fairly complete 
picture of its development from egg to adult. 

Genus MASTURUS Gill 
Masturus Gill, 1884, Proc. U.S. Nat. Mus. 7: 425. Type: Orthagoriscus oxyuropterus Bleeker. 

The study of this genus has been greatly facilitated by the careful bibliographical 
work of Gudger, who studied the records over a number of years, added several new 
ones, and finally in 1937 published a work dealing with the structure of the caudal 
region and another summarizing the knowledge of the genus up to that date. The 
latter two works are of great value, and the remarks which I make in the pages which 
follow are based largely upon them and should be considered with constant reference 
to them. 

The distinctness of Masturus from Mola had already been acknowledged by Steen- 
strup & Lutken (1898) , and discrimination between the post-larval forms was achieved 
by Schmidt (1921). The secondary post-larval stage of Masturus is characterized by 
enormous elongation of the 'cornicles' (Fig. 6). But it remained for Gudger to dis- 
entangle the confusion in the literature, and it is no doubt because these necessitated 
a chronological arrangement of his data that he was unable to recognize the two 
forms involved. But an analvsis of the records leaves little room for doubt that there 



THE OCEAN SUNFISHES (FAMILY MOLIDAE) 99 

are indeed two forms, one the generally accepted M. lanceolatus (Lienard), the other 
apparently taking M. oxyuropterus (Bleeker) as the earliest name. These will be 
diagnosed on a later page, but it is necessary to note their existence at this point in 
order to clarify the discussion which follows. I must stress now, however, that they 
are regarded here as species only because we have no knowledge to the contrary, 
but I suspect that they may prove to be the sexes of one species. Not one of the 




A, 



Fig. 6. Post-larvae of Masturus. 
Ostracion hoops' stage (2-8 mm.) ; B, ' Molacanthus 



(5 mm.). 



mm.) 
(After Schmidt.) 



stage 



specimens so far recorded has been sexed. Raven, the only person to make a dissec- 
tion, does not even mention the gonads. 

Masturus is peculiar among the Molidae in the possession of a pronounced lobe a 
little above the centre of the clavus. Gudger continually stressed the dorsal situation 
of this lobe, apparently as evidence that it could not be the remains of the larval tail ; 
this is not a very good point, for his own anatomical figures show that the lobe is 
associated with the end of the vertebral column. In other Plectognathi the vertebral 
column lies dorsally until it enters the caudal peduncle, where it lies approximately 
in the central long axis of the body. The fact that in the Molidae the vertebral column 
is dorsally placed at its hind end is therefore interesting as a further demonstration 
that the posterior part of the column is lost. 

It is my belief that the lobe on the clavus of Masturus can truly be called the 



ioo THE OCEAN SUNFISHES (FAMILY MOLIDAE) 

'caudal lobe ', for all the illustrations of its anatomy so far given seem to demonstrate 
that the slender rays supporting it are caudal rays. The first to be published was that 
by Ryder, after a drawing by Putnam; it was reproduced by Gudger, and is now 
copied as my Fig. 7 A. It is interesting in that the dorsal and anal rays of the clavus 
are shown branched, a feature shown only once elsewhere in the literature (Gudger, 
1939), perhaps because the tips are so often broken off in young specimens. They are 




Fig. 7. Caudal skeletons of A, Masturus oxyuropterus, copied from Ryder, 1886; B, Masturus 

lanceolatus, copied from Gudger, 1937. 
c, caudal rays; inn, interneural bones; inh, interhaemal bones. 

thus distinguishable at a glance from the simple caudal rays in the middle, but it is 
probable that the outermost two of the latter are also of dorsal and anal origin, for 
they have each a small skeletal support. 

The interspinous bones supporting the clavus are shown completely fused with the 
hindmost remaining haemal spine. Comparison with other dissections shows that 
these must in fact have been distinct elements. The shape of the supporting bones of 
the dorsal and anal fin lobes is obscured by the inclinator muscles in this figure, but 
the drawing of these muscles is interesting in helping to show their character after the 
lateralis mass has been removed. 

Each of the rays in the clavus is supported by an interspinous bone, with the excep- 
tion of the middle four ; these are associated with the last of the remaining vertebrae, 



THE OCEAN SUNFISHES (FAMILY MOLIDAE) 101 

which has no neural or haemal elements. There is no apparent reason why, if they 
also are dorsal and anal rays, they should not have their supports ; but if they are 
caudal rays they cannot be expected to be borne on hypurals, since these and other 
posterior vertebral elements have been lost. The presence of only four of these un- 
supported rays and the equal length of the dorsal and anal fin bases shows that 
Putnam's fish was a Masturus oxyuropterus ; two other dissections of this form have 
been illustrated — that byGudger (1937 a, p. 41, fig. 27), and that by Raven (19396). 
The first does not show the internal skeleton, and one of the caudal rays is doubled, 
as can be seen by the nature of its basal cartilage ; but Raven's illustration, drawn by 
Helen Ziska, is admirable, and agrees in all essentials with that by Putnam. The only 
illustration showing the anatomy of M. lanceolatus is that given by Gudger in the 
work just quoted, based upon a young specimen (the same size as Putnam's) which 
was stained with alizarin and cleared. During the staining process some of the 
elements, notably the interneural supports of the clavus, were displaced, but I am 
satisfied that nothing was lost. This illustration is copied here as my Fig. 7 b. Here 
it will be seen that the central lobe of the clavus is supported by eight rays whose 
only skeletal support is the last vertebra (which has no neural or haemal elements) . 
Above these are five rays which can be associated with the five interneural bones 
which have been displaced from the horizontal during preparation. Below them are 
nine rays which belong to nine interhaemals, the lower three of which have been dis- 
placed forward. The presence of eight rays in the caudal lobe of the clavus and the 
greater length of the base of the dorsal fin lobe as compared with that of the anal fin 
lobe shows this to have been a specimen of Masturus lanceolatus as identified by 
Klunzinger (a figure of whose specimen is given by Gudger). 

It is admittedly hazardous to speak of caudal rays when the hypural bones are lost, 
since in normal fishes caudal rays are distinguishable only by their association with 
the hypurals. But I feel convinced that these central rays of the clavus in Masturus 
are homologous with the hypocaudal rays of the more generalized forms, and it 
remains for me to suggest how it is possible for them to persist although their skeletal 
supports are lost. 

It has to be borne in mind that two opposing forces are involved during the de- 
velopment of the caudal region, interacting in different proportion at successive stages. 
First there is the reduction of the larval tail and the atrophy of the posterior vertebral 
elements, and secondly the normal growth of body and fins. 

The first process evidently begins at an early age, for Schmidt has figured a larval 
specimen in which, as Gudger has pointed out, dorsal and anal rays are present but 
not caudal rays ; development of the latter is retarded. To see how this fact may affect 
later stages it is necessary to consider what occurs in the Triacanthodidae, the most 
primitive family of Plectognathi. In the larval Triacanthodid (Fig. 8), the caudal 
rays are twelve in number, as in most Plectognathi, but the last four lie in relation 
to the end of the notochord, which will later become ossified as the urostyle; the 
anterior eight belong to the last few myotomes. Degeneration of the tail from the 
rear will mean that the end of the notochord is lost first, and if this occurs before the 
hypocaudal rays appear not more than eight of them will develop. The eight slender 
rays of Masturus lanceolatus thus become intelligible and significant, while the 



102 THE OCEAN SUNFISHES (FAMILY MOLIDAE) 

presence of only four in M. oxyuropterus suggests that reduction has proceeded still 
farther before caudal rays begin to develop in this form. Comparison with the larval 
Mola figured by Sanzo (1939) is interesting in this connexion, for it will be seen that 
if in his specimen hypocaudal rays were developed, they would not be associated 
with myotomes, and this probably accounts for their absence in that genus. 

As the caudal rays become stronger the axial structures decrease rapidly, so that 
by the time the rays are brought to the homocercal position there are no hypurals for 
their support, nor neural or haemal elements for the last few vertebrae ; but normal 




Fig. 8. Caudal region of post-larval Triacanthodid fish, showing relationship 
of hypocaudal rays to notochord and myotomes. 



body growth has extended the posterior parts of the dorsal and anal fins with their 
supporting structures backward and downward to fill the void. This process is 
probably correctly demonstrated by Ryder's diagram, upon which mine is based 
(Fig. 1) , in which the region of atrophy is delineated by the broken line. The vertebrae 
with their neural and haemal arches and spines are lost, but the interneural and 
interhaemal spines develop in relation to the fins in the normal manner except that 
they ultimately become tilted nearly at right angles to the last developed vertebral 
elements (Fig. 7). The number of these interspinous bones does not give a reliable 
estimate of the number of vertebrae that have been lost, because reference to the 
dissections shows that more than one may be associated with each neural or haemal 
spine, while of course the last few vertebrae are probably not associated with inter- 
spinous bones at all. Ryder, of course, thought Putnam's young fish was a Mola 
and that the caudal rays were completely lost in the adult. A curious feature of the 
posterior migration of the dorsal and anal fins is that, while in the lobes the rays are 
more numerous than the interspinous bones, each of those in the clavus has its own 
supporting element ; this might be taken to indicate that the central rays, which I 



THE OCEAN SUNFISHES (FAMILY MOLIDAE) 103 

regard as caudal, are simply the last dorsal and anal rays, which are therefore more 
numerous than their supports in this region also, but there seems no good reason why 
the odd rays should all be crowded at the end. A more difficult argument to combat 
is that the supporting elements of these last rays cannot develop because of the 
presence of the vertebral column ; against this I can only point out that in Putnam's 
fish two elements lie behind the last vertebra, and there seems no reason why, if the 
odd four rays were in the same series, their supporting bones should not be there also. 
In fact, the presence of the two elements mentioned is reminiscent of the condition 





INT 



Fig. 9. Masturus oxyuropterus , late post-larva (21 mm.), in British Museum collection. B, dis- 
section of same specimen, showing presence of air-bladder. 
ab, air-bladder; h, heart; k, kidney; l, liver; int, intestine; spl, spleen. 



shown in Cyema atrum by Trewavas (1933), who identified the two small ossifications 
as hypurals. But since the last caudal vertebrae are so obviously lost in the Molidae 
it would be incautious to speak of hypurals here. 

McCulloch has left us drawings of very young examples of both M. lanceolatus and 
M. oxyuropterus, at the stage when the larval tail is not quite lost, the small peduncle 
bearing its allotted quota of caudal rays and the dorsal and anal fins extending round 
to meet them. Knowing what a careful observer and excellent draughtsman McCul- 
loch was, I am prepared to accept these as good evidence. Eventually, at the better 
known stage of 50 mm. or thereabouts, there is no sign of the original tail, but the 
caudal rays project beyond the rest of the clavus as the basis for the ultimate central 
lobe. Gudger believed that even these central rays were lost, at what he called the 
'square-tailed ' stage, but as this was based on the two obviously damaged specimens 
of Steenstrup & Liitken, this seems to be improbable — a point which Dr. Gudger 
himself has conceded in a letter to me. 

As a matter of interest I may mention here that in these small specimens it appears 
that the air-bladder is still present ; one which I dissected (Fig. 9 b) had a very 

zool. 1. 6 N 



104 



THE OCEAN SUNFISHES (FAMILY MOLIDAE) 



delicate, bubble-like structure at the centre of mass, which unfortunately collapsed 
while I was examining it. At this planktonic stage in its development such an organ 
is not surprising, and of course the Molidae are evolved from fishes in which the air- 
bladder is well developed, but it is worth noting that the statement that an air-vessel 
is absent in this family is probably true only of adults. 




Fig. io. Masturus lanceolatus, adult. Singapore. (After Smedley, 1932.) 

With the development of the skeletal structures (poorly ossified though they are) 
atrophy proceeds no farther, and the processes of growth produce what later changes 
we can observe in the fish. An extension of the dermis and its collagenous sub- 
stratum, probably that which would develop over the caudal peduncle in a more 
normal fish, eventually covers the caudal lobe and the whole clavus. 

These are what seem to be the main features in the development of the clavus of 
Masturus, but there is a certain amount of individual variation. In M. lanceolatus 
the presence of eight caudal rays seems to be fairly consistent, but the middle ones 
are sometimes represented only distally — whether their proximal ends atrophy in the 
early stages or degenerate later is not evident. In M. oxyuropterus four caudal rays 



THE OCEAN SUNFISHES (FAMILY MOLIDAE) 105 

are usual, but may occasionally be five (as in a dissection figured by Gudger, wherein 
one of the rays had split or doubled as shown by its supporting cartilage) or rarely 
three. On published evidence the number of rays supporting dorsal fin, clavus, and 
anal fin respectively appear to differ very greatly, but most of these are of doubtful 
value, for an accurate count can only be made by dissection (except possibly in 
stuffed specimens) . As an example of this may be quoted the description by Gudger, 
a careful worker, of the specimen he obtained for the American Museum of Natural 
History. In this he counted, on external examination, 'D. + C.+A. complex = 60' ; 
the fish is in other respects M. oxyuropterus, so that this high count would cast doubt 
on its distinctness from M. lanceolatus. But later Raven dissected this same fish, 
and his illustration shows distinctly the total of fifty-five rays which is usual in 
M. oxyuropterus. Consequently it has not seemed expedient to give any definite 
statement of the number of rays to be found in dorsal and anal fins and clavus 
respectively, but only to indicate the total number, which seems to be characteristic 
for each species. 

Whether or not I am correct in calling them caudal rays, the presence of median 
rays unsupported by interspinous bones is characteristic of Maslurus. In the adults 
all the rays of the clavus are simple, without distal ossifications. There is always a 
median projection to the clavus, and the body is rather more elongate than that of 
Mola, especially in the early stages. Osseous tubercles, the remains of post-larval 
spines, seem never to be retained anywhere on the body of the adult. 

Two forms can be recognized, treated here as species, but I suspect that further 
study will show them to be the sexes of one. They have been taken in the same 
localities, and sometimes together. The sexual dimorphism found to be present in 
Mola mola (p. 117) lends support to this idea. But with no knowledge of the sex of 
any recorded individual I can but state their characteristics and apply available 
names to them pending further information. 

Since all the literature before 1939 has been fully quoted and discussed by Gudger, 
I have not thought it necessary to repeat it all below, particularly as a number of 
records cannot be assigned with certainty, but full and discriminating reference has 
been made to Gudger's papers. 

KEY TO THE SPECIES OF MASTURUS 
I. Profile of lower jaw more convex, usually projecting beyond the upper. Upper 
profile of head evenly convex. Base of dorsal lobe conspicuously longer than that 
of anal fin. Dorsal and anal fins and clavus with a total of 60 to 62 rays. Caudal 
lobe of clavus 1 longer than head in perfect specimens (often mutilated), sup- 
ported by eight (rarely 7 or 9) rays . . . . .1. lanceolatus 
II. Profile of lower jaw less convex, straight or concave, not projecting beyond the 
upper. Upper profile of head with distinct concavity above the eyes. Bases of 
dorsal and anal fin lobes about equal. Dorsal and anal fins and clavus with a total 
of 55 to 57 rays. Caudal lobe of clavus shorter than head, supported by 4 (rarely 
3 or 5) rays . . . . . . . . 2. oxyuropterus 

The uncertainty of authors as to where dorsal and anal fins end and clavus begins, 

1 Measured from ' hinge ' of clavus to tip. 



106 THE OCEAN SUNFISHES (FAMILY MOLIDAE) 

and the obvious inaccuracy (already mentioned) of fin-ray counts made upon 
external examination, militates against giving counts for individual fins, but it may 
be, as suggested by Gudger's cleared specimen, that in M. lanceolatus there are more 




Fig. ii. Masturus oxyuropterus, adult. Tahiti. (After Gudger, I935-) 



rays in the dorsal lobe than in M. oxyuropterus. In the latter, on the other hand, the 
number of claval rays supported on interhaemal bones seems to be greater (10 to 12) 
than in M. lanceolatus (9). 

Although usually these types seem to be recognizable at an early age, there are 
some doubtful cases among young specimens, as might be expected if they were the 



THE OCEAN SUNFISHES (FAMILY MOLIDAE) 107 

sexes of one species. For example, if M. lanceolatus should be the male, it might be 
more like the female (M. oxyuropterus) when young, as in many other fishes, and in 
fact small examples of the oxyuropterus type seem to be the more common. 

Masturus lanceolatus (Lienard) 

Orthagoriscus lanceolatus Lienard, 1840, Revue Z00L: 291; 1841, Magasin Zool. (2) 3 (Poiss.) : 

pi. 4. 
Orthagoriscus mola Khmzinger, 1871, Verh. ZooL-Bot. Ges. Wien, 21: 648; Gunther, 1880, Introd. 

Stud. Fish.: 175, fig. 94; Perugia, 1881, Ann. Mus. Stor. Nat. Genova 27: 365, fig. 
Mola mola Collett, 1896, Result. Camp. Sci. Monaco 10: 163, pi. 6, fig. 1. 
Ranzania truncata Steenstrup & Lutken, 1898, K. danske vidensk. Selsk. Skr. (6) 9: pi. 6, fig. C. 

(Not of Jordan & Gilbert, 1883.) 
Mola (Molacanthus) sp. McCulloch, 1912, Proc. Linn. Soc. N.S.W. 37 (3) : 553, pi. 58. 
Mola lanceolata Schmidt, 1921, Medd. Komm. Havundersog. Kbh., Fisk. 6 (6): pi. 1, figs. 4, 5; 

1932, Dana's Togtomkr. Jord.: 255, fig. 167 (part.); Barnard, 1927, Ann. S. Afr. Mus. 21: 

987, fig. 31 ; Ehrenbaum, 1936, Handb. Seefisch. Nordeurop. 2: 88; J. L. B. Smith, Sea Fish. 

5. Afr. : 422, fig. 1214. 
Masturus lanceolatus Hubbs & Giovannoli, 1931, Copeia, 1931: 135; Gudger, 1935, Amer. Mus. 

Novit. 778: 1, fig. 1 ; Gudger & McDonald, 1935, Sci. Mon. 41: 1, figs. 4-9, 11, 14, 15 ; Rivero, 

1936, Amer. Nat. 70: 92, fig.; Palmer, 1936, Science, 83: 597; Gudger, 1937, Ann. Mag. Nat. 

Hist. (10) 19: 9, fig. 6; 15, fig. 10; 31, fig. 19; 33, fig. 20; 34, fig. 21 ; 38, fig. 23; Proc. Zool. Soc. 

Lond. 107 (A) (3) : 353 (part.), text-figs. 1, 2, 4, 6, 7, 8, 9, 14 ( ?), 16, 20, 21 ( ?) ; pi. 1, figs. 3, 4 ; 

pi. 2, figs. 5, 6 ; pi. 4, fig. 10 ; 1939, /. Elisha Mitchell Sci. Soc. 55 (2) : 305 ; Brimley, 1939, ibid. 

295, pi. 28; Fitch, 1950, Calif. Fish Game, 36 (2) : 65. 

Lienard 's figure cannot be said to be notable for its faithful representation, but as 
it shows the caudal lobe indubitably much longer than the head and the base of the 
dorsal fin longer than that of the anal, it shows to which of our two forms the name is 
applicable. The fin-rays are always more easily seen in dried specimens of these 
fishes, and so Klunzinger's stuffed example shows the structure of the clavus very 
well ; it is closely similar (in this 65-in. example) to that of Gudger's 53-mm. cleared 
specimen. Other figures in Gudger's papers which appear to represent this form are 
stated in the above synonymy. Where the caudal lobe is mutilated or otherwise 
doubtful the broad dorsal base and rather pugnacious-looking 'chin' are the most 
useful distinguishing characters. 

It grows to a great size, the largest recorded specimen being 10 ft. long and 11 ft. 
3 in. from the tip of dorsal to tip of anal fins. In our collection it is represented 
only by the post-larval specimen figured by Gunther. 

Recognizable records of adults are from the Atlantic, off Florida, Havana, North 
Carolina, and Table Bay, South Africa, from the Red Sea, and from the Pacific at 
Tahiti. Young specimens have been taken off Alabama, Teneriffe, and in the South 
Seas. As this paper goes to press Fitch (1950) states that 100 post-larvae J to 2 in. 
in length have been taken from the stomachs of tuna in Hawaiian waters. 

Masturus oxyuropterus (Bleeker) 

Orthagoriscus spinosus Gatchet, 1832, Act. Soc. Linn. Bordeaux 5: 253. (Not of Cuvier, 181 7.) 
Orthagoriscus oxyuropterus Bleeker, 1873, Versl. Akad. Amst. (2) 7: 151, fig. 



io8 



THE OCEAN SUNFISHES (FAMILY MOLIDAE) 



Mola rotunda Ryder, 1886, Rep. U.S. Fish. Comm. (1884): 1027, pi. 8, fig. 5. (Not of Cuvier, 

1798.) 
Ranzania truncata Steenstrup & Lutken, 1898, K. danske vidensk. Selsk. Skr. (6) 9 (1) : 98, pi. 6, 

figs. D, E. (Not of Jordan & Gilbert, 1883.) 
Mola (Molacanthus) sp. McCulloch, 1912, Proc. Linn. Soc. N.S.W. 38 (3): 553 {part.), pi. 59. 



FLORIDA _ 


9; 




FLORIDA 








8 : 




ATLANTIC 

FLORIDA ^ 


7; 




N. CAROLINA--'' 

TABLE BAY 

FLORIDA -' 






RED SEA 


6 




HAVANA 








5 




TAHITI . 


4 


: 




3 


:. _, 




2 





MIAMI , FLA 



HAWAII 
SINGAPORE 



FLORIDA 
AMBOINA 



N CAROLINA 



FEET 
and inches 



Fig. 12. Diagram showing comparative ranges of size for the two species 
of Masturus, based on recognizable records of adult specimens. 



Mola mola Townsend 1918, Bull. N.Y. Zool. Soc. 21: fig. (not of Linnaeus) ; Collett, 1896, Result. 

Camp. Set. Monaco, 10: 163 (part.) pi. 6, fig. 1. [*& 

Mola lanceolata Schmidt, 1921, Medd. Komm. Havundersog. Kbh. Fisk. 6 (6) {part.): pi. 1, fig. 

6; Smedley, 1932, Bull. Raffles Mus. 7: 17, pi. 
Masturus lanceolatus Jordan & Jordan, 1925, Mem. Carneg. Mus. 10: 89, fig. 7; Gudger & 

McDonald, 1935, Sci. Mon. 41: 1, figs. 3, 10, 12, 13 ; Gudger, 1935, Copeia, 1935: 35, figs. 1, 2 ; 

1937, Ann. Mag. Nat. Hist. (10) 19: 1 (part.), text-figs. 18, 22, 26, 27; 1937, Proc. Zool. Soc. 

Lond. 107 (A) (3): 353 (part.), text-figs. 5, 10, 12, 13, 15, 18 (?), 19, 22, pi. 1, figs. 1, 2, pi. 3, 

n g- 9 ( ?), pi. 4, fig. ii, pi. 5, fig. 17; 1939, /. Elisha Mitchell Sci. Soc. 15 (2) : 305 (part.), figs. 



THE OCEAN SUNFISHES (FAMILY MOLIDAE) 109 

1-5 ; Brimley, 1939, ibid. 300, pi. 29 ; Raven, 1939, Bull. Amer. Mus. Nat. Hist. 76 (4) : 143, pi. 
2; Hardenberg, 1939, Treubia 17 (2): 121 ; Clark, 1949, Amer. Mus. Novit. 1397: 7, fig. 9. 

A high proportion of the young specimens recorded seem to belong to this form, 
but a possible explanation of this is given on page 107. The small number of support- 
ing rays in the caudal lobe, the equal bases of dorsal and anal fins, and the compara- 
tively weak-looking 'chin' are recognizable even in McCulloch's 10-mm. specimen. 
The concavity of the dorsal profile of the head, however, is not noticeable in very 
small specimens, but it is already apparent in the 152-mm. example figured by 
Gudger (1939). The latter paper is also interesting in that it shows branching at the 
tips of the rays of the clavus, like that illustrated by Ryder (Fig. 7 a in this paper), 
but very much smaller, evidently in process of reduction. Only these two records of 
such branching exist, probably because the tips of the rays have been damaged in 
most small specimens, and the branching is lost with age. 

This form is so often taken in the same locality as the preceding that it is almost 
certainly a sex of that species ; in some instances young specimens of both forms have 
been taken from a single predatory fish (e.g. McCulloch (1912), whose 13-mm. specimen 
is M.lanceolatus and his 10-mm. specimen M. oxyuropterus ; or Gudger (1939), whose 
125-mm. fish is M. oxyuropterus, whereas at least the 127-mm. fish, and possibly the 
130-mm. specimen also appears to be M. lanceolatus) . 

It will be noticed that in each case the M. oxyuropterus is slightly the smaller, and 
the records of this form do tend to lie about a lower range of size (Fig. 12). The 
largest record seems to be the 'Miami Masturus no. Ill' of Gudger, figured by 
Gudger & McDonald, though the identification of this badly slung specimen is a little 
doubtful. It was 7 ft. in length. 

Recognizable records of adults of this form are from the Atlantic at North Carolina 
and Florida, from Singapore and Amboina, and from the Pacific at Hawaii. Young 
specimens have been taken at Florida, the Sargasso Sea, the Azores, and in the South 
Pacific. 

A young specimen (Fig. 9) of unknown provenance is in our collection, and a plaster 
cast of the specimen dissected by Raven is exhibited in the fish gallery of the British 
Museum (Natural History). 

Genus MOLA Koelreuter 

Mola Koelreuter, 1770, Novi Comment. Acad. Petropol. 8: 337. Type: Mola aculeata Koelreuter 

(= Tetraodon mola Linnaeus, young). 
Orthragoriscus Bloch & Schneider, 1801, Syst. Ichth.: 510. Type: Tetraodon mola Linnaeus. 
Cephalus Shaw, 1804, Gen. Zool. 5: 437. Type: Tetraodon mola Linnaeus. 
Orthragus Rafinesque, 1810, Caratt. Sicilia: 17. Type: Orthragus luna Rafinesque (= Tetraodon 

mola Linnaeus). 
Diplanchias Rafinesque, 1810, ibid. Type: Diplanchias nasus Rafinesque. 
Orthagoriscus Cuvier, 1817, Regne Anim., ed. 1, 2: 149. Type: Tetraodon mola Linnaeus. 
Pedalion Swainson, 1839, Nat. Hist. Fish. 1: 199. Type: Pedalion gigas (Guilding) Swainson. 
Molacanthus Swainson, 1839, ibid. 2: 329. Type: Molacanthus pallasi Swainson (= Tetraodon 

mola Linnaeus). 
Ozodura Ranzani, 1839, Novi Comment, acad. Sci. Inst. Bonon. 3: 80. Type: Ozodura orsini 

Ranzani. 
Tympanomium Ranzani, 1839, ibid., table. Type: Tympanomium planci Ranzani. 



no THE OCEAN SUNFISHES (FAMILY MOLIDAE) 

Trematopsis Ranzani, 1839, ibid., table. Type: Trematopsis willoughbii Ranzani. 

Pallasina Nardo, 1840, Ann. Sci. Regno Lombardo-V eneto 10: 10, 112. Type: Pallasina pallasi 

Nardo (larval form). 
Acanthosoma De Kay, 1842, Nat. Hist. New York (Zool.) 3: 330. Type: Acanthosoma carinatum 

De Kay (= Tetraodon mola Linnaeus, young). 
Aledon Castelnau, 1861, Mem. Poiss. Afr. austr.: 75. Type: Aledon storeri Castelnau. 

Closely related to Masturus, but differing in that the clavus is supported entirely 
by elements from the dorsal and anal fins. The form of the body is relatively shorter, 
conspicuously so in the young, and the post-larval spines are not entirely lost, the 
base of one at the chin or one on the snout, or both, remaining as a low bony boss in 
the largest examples. 

Very few post-larval specimens of Mola have been found, but the smallest, 5 mm. 
long, shows that there is an ' Ostracion hoops ' stage, and several examples of the 
secondary post-larval or 'Molacanthus' stage have been described ; it is not known 
whether the 'cornicles' are ever as long as those of Masturus. 

Although a number of naturalists have believed in the existence of several species 
of Mola, and Ranzani went so far as to recognize five genera and eleven species, it has 
generally been believed, especially during this century, that only one widely dis- 
tributed species is admissible. 

My studies, however, show that while Mola mola is indeed wide-ranging, it is 
largely or entirely replaced in the South Pacific by a second species, distinguishable 
as follows : 

KEY TO THE SPECIES OF MOLA 

I. Clavus supported by about 16 rays, 12 of which bear ossicles; the ossicles much 
broader than the spaces between them, and forming the margin of the clavus ; 
those borne on paraxial rays separate, much smaller than the others. No band 
of reduced denticles between dorsal and anal fins . . . 1. ramsayi 

II. Clavus supported by about 12 rays, 8 or 9 of which bear ossicles; the ossicles 
widely separated, invested with cuticle, which grows beyond them to form lobes 
in large examples ; those borne on paraxial rays united to form a single ossicle 
larger than all the others. A band of reduced denticles, smoother to the touch, at 
base of clavus from dorsal to anal fin . . . . .2. mola. 

The term 'paraxial rays' refers to the pair of supporting rays of the clavus the 
proximal ends of which lie nearest to the end of the vertebral column. The smooth 
band between dorsal and anal fins in M. mola is usually visible, marked by a fold 
posteriorly, and often differently coloured from the rest of the fish ; in doubtful cases 
the tips of the fingers will discern that this area is less rough than the body in front 
of it and the clavus behind it. 

Mola ramsayi (Giglioli) 

Orthagoriscus truncatus Hutton, 1872, Fish. New Zealand: 73. (Not of Fleming, 1828.) 
Orthagoriscus mola Castelnau, 1872, Proc. Zool. Acclim. Soc. Vict. 1: 211; 1875, Res. Fish. 

Austral.: 3; Hutton, 1873, Trans. Proc. N.Z. Inst. 5: 271; Macleay, 1875, Proc. Linn. Soc. 

N.S.W. 1: 12; Johnston, 1883, Pap. Roy. Soc. Tasm.: 137; 1891, ibid.: 38; Hamilton, 1886, 



THE OCEAN SUNFISHES (FAMILY MOLIDAE) in 

Trans. Proc. N.Z. Inst. 18: 135; Williams, 1893, ibid. 25: no, pi. 8a; Drew, 1897, ibid. 
29: 286 ; Parker, 1897, ibid. : 627 ; ? Fletcher, 1929, Proc. Linn. Soc. N.S.W. 54: 225, 227. (Not 
of Cuvier, 181 7.) 



Fig. 13. Mola ramsayi, adult, 2130 mm. long, New South Wales. (Drawn from 
the type of the species in the British Museum collection.) 

Orthragoriscus ramsayi Giglioli, 1883, Nature, Lond. 28: 315; Ramsay, 1883, Cat. N.S.W. Court. 

Intern. Fish. Exhib.: 43. 
? Orthagoriscus eurypterus Philippi, 1893, Chilen. Fische: 15, pi. 6, fig. 1 (not seen). 
Mola mola Waite, 1907, Rec. Canterbury [N.Z.] Mus. 1: 34; 1913, Trans. N.Z. Inst. 45: 223, 

pi. 9 ; 1921, Rec. S. Aust. Mus. 2: 198, fig. 332 ; 1923, Fish. S. Austral. : 230, fig. ; Phillips, 1919, 

ZOOL. I. 6 O 



2 THE OCEAN SUNFISHES (FAMILY MOLIDAE) 

Rep. Dom. Mus. N.Z.: 6; 1926, N.Z. J. Sci. Tech. 8 (3): 169, figs. 1-3; McCulloch, 1922, 
Aust. Zool. 2 (3): 130, fig. 374 a; 1930, Mem. Aust. Mus. 5 (3): 436 (part.) ; Schneider, 1930, 







Fig. 14. Mola ramsayi, young adult, 410 mm. long, South Australia ( ?). (From 

specimen in spirits in the British Museum collection.) 

po, paraxial ossicles. 

Rev. Chil. Hist. Nat. 34: 200, figs. 36, 37 ; Fowler, 1945, ibid. 45-47: 170, fig. ; Morrow & Mauro 
1950, Copeia, 1950: 108, fig. 4 c. 
Mola ramsayi Whitley, 1931, Rec. Aust. Mus. 18 (3) : 126 (part.), pi. 16, figs. 3, 4. 



THE OCEAN SUNFISHES (FAMILY MOLIDAE) 113 

All the New Zealand records, most of the Australian, and the few Chilean specimens 
appear to belong to this species, though in many cases it is not possible to be certain. 
It may be assumed, therefore, that in the South Pacific it replaces the wide-ranging 
M. mola. The two meet, however, in the Australian area, for Stead, McCulloch, and 
Whitley have all figured specimens which were undoubtedly M. mola, Whitley 
including his specimen with one of the true M. ramsayi in the same paper under the 
latter name. 

The type of Orthragoriscus ramsayi Giglioli is in the British Museum (Nat. Hist.). 
Its locality was given as 'Southern Hemisphere', but a label accompanying the 
specimen states 'New South Wales', and it is known to have been taken on that 
coast (fide Whitley, 1931). It was exhibited at the International Fisheries Exhibition 
in London in 1883 and later presented to the Museum by the Commissioners of the 
Exhibition. It is a very large stuffed skin, now in a rather dilapidated condition. 
The total length is 213 cm. (6 ft. 8 in.). 

We have, fortunately, a second specimen, in spirits — much smaller, of course ; it 
is without a definite locality, but almost certainly from South Australia, since it was 
in a collection of specimens presented by the Zoological Society, several of which 
were typical South Australian species and all of which would be likely to occur there. 
It agrees very well with the excellent figure given by Waite (1923), and removes any 
doubt as to the distinctness of the species from M. mola. 

The type is not by any means the largest recorded specimen of M. ramsayi. That 
distinction apparently goes to one taken on 12 December 1889 in Poverty Bay, and 
recorded by Williams as measuring 9 ft. 8 in. and weighing 3^ tons. 



Mola mola (Linnaeus) 

Tetraodon mola Linnaeus, 1758, Syst. Nat. ed. 10, 1: 334; Pennant, 1776, Brit. Zool. 3: 131, 

pi.; Migliorini Spinola, 1843, Poiss. Genes: 14. 
Tetrodon mola Briinnich, 1768, Ichth. massil.: 8; Gmelin, 1778, Syst. Nat. Linn.: 1447; Retzius, 

1785, K. Svensk. Vetensk. Akad. Handl. 6: 115; Bonnaterre, 1788, Tabl. Encycl. Meth.: 25, 

pi. 17, fig. 54; Lacepede, 1798, Hist. Nat. Poiss. 1: 509; Retzius, 1800, Fauna Suec: 310; 

Donovan, 1803, Nat. Hist. Brit. Fish. 2: pi xxv. 
Mola aculeata Koelreuter, 1770, Novi Comment. Acad. Petropol. 8: 337. 
Diodon mola Pallas, 1777, Naturgesch. Thieve 8: 41, pi. 4, fig. 7; Bloch, 1785, Naturgesch. 

ausldnd. Fische 1: 75, pi. 128; Jacob. 1826, Dublin Phil. J. 2: 443, pi. 
Mola rotunda Cuvier, 1798, Tabl. Elem. Nat. Hist. : 323 ; Jordan, 1881, Proc. U.S. Nat. Mus. : 70 ; 

Jordan & Gilbert, 1883, Bull. U.S. Nat. Mus. 16: 865; Petersen, 1884, Vidensk. Medd. naturh. 

Foren. Kbh.: 159; Smith, 1885, W. Amer. Sci. 1 (7): 45; Linton, 1897, Proc. U.S. Nat. Mus. 

19: 788, 812, 824; Steenstrup & Liitken, 1898, K. Danske vidensk. Selsk. Skr. (6) 9 (1) : 28, 

pi. 1 ; Murray & Hjort, 1912, Depths of the Ocean: 119, 607, 615, 697, figs. 102, 507; Schmidt, 

1921, Medd. Komm. Havundersog. Kbh. Fisk. 6: 1, figs. 1, 5, 6, 10 b, 12, pi. 1, figs. 1, 2 ; 1926, 

Nature, Lond. 117: 80, figs. 1,2; Ehrenbaum, 1936, Handb. Seefisch. Nordeurop. 2: 86, fig. 68 ; 

Jensen, 1940, Vidensk. Medd. nat. Foren. Kbh. 104: 319. 
Orthragoriscus mola Bloch & Schneider, 1801, Syst. Ichth.: 510; Turner, 1862, Nat. Hist. Rev.: 

185, pi. 6, figs. 4-6; Beneden, 1871, Mem. Acad. R. Belg. 38; Jeude, 1890, Notes Leyden Mus. 

12: 189, pi. ; Roon & Pelkwijk, 1939, Zool. Meded. Leiden 22: 65, figs. 1, 2. 
Orthragoriscus fasciatus Bloch & Schneider, 1801, Syst. Ichth.: 511. 

ZOOL. I. 6 2 



ii 4 THE OCEAN SUNFISHES (FAMILY MOLIDAE) 

Orthragoriscus hispidus Bloch & Schneider, 1801, ibid.: 511. 

Cephalus brevis Shaw, 1804, Gen. Zool. 5: 437, pi. 175; Neill, 1811, Mem. Werner. Soc. 1: 546; 
Mitchill, 1815, Trans, lit. phil. Soc. N.Y. 1: 471 ; Swainson, 1839, Nat. Hist. Fish. 1: 199. 

Cephalus pallasianus Shaw, 1804, Gen. Zool. 5: 440. 

Orthragus luna Rafmesque, 1810, Caratt. Sicilia: 17-18; Indice Siciliana: 40. 

Diplanchias mola Rafmesque, 1810, ibid. 

Cephalus mola Risso, 1810, Ichth. Nice: 60; Poey, 1868, Repert. Cuba 2: 433. 

Orthagoriscus mola Cuvier, 1817, Regne Anim., ed. 1, 2: 149; Fleming, 1828, Hist. Brit. Anim.: 
175; Nilsson, 1832, Prodr. Ichth. Scandinav.: 111; Jenyns, 1835, Man. Brit. Vertebr. Anim.: 
490; Storer, 1839, Fish. Massachusetts: 170, pi. 3, fig. 1; Swainson, 1839, Nat. Hist. Fish. 2: 
329, fig. 107; Bellingham, 1840, Mag. Nat. Hist, (n.s.), 4: 235; Bennett, 1840, Narr. Whaling 
Voy. 2: 262 ; Wellenbergh, 1840, Dissert. Inaug., Lugd. Batav., pi. ; Goodsir, 1841, New Philos. 
J. 30: 188, pi. 4; De Kay, 1842, Nat. Hist. N.Y. {Zool.), 3: 331, pi. 59, fig. 193; Storer, 1846, 
Mem. Amer. Acad. Arts Sci. n.s. 2: 495; Dilwyn, 1848, Mater. Fauna Swansea: 15; Parlby, 
1848, Proc. Zool. Soc. Lond. 17: 6; 1850, Ann. Mag. Nat. Hist. (2) 5: 53 ; Schlegel, 1850, Fauna 
Japonica (Poiss.): 288, pi. 127; Costa, 1850, Fauna Regn. Napoli (Pesci, Plettognathi) : 28, 
pis. 63-64; Smith, 1851, Ann. Mag. Nat. Hist. (2) 8: 347; Kroyer, 1852, Danmarks Fisk. 
3: 732; Embleton, 1854, Trans. Tyneside Nat. 2: no, pi. 3 ; Nilsson, 1855, Skandinav. Fauna: 
697 ; Thompson, 1856, Nat. Hist. Ireland 4: 243 ; Kolliker, i860, Verh. phys-med. Ges. Wurzburg 
10: xxxviii; Cleland, 1862, Nat. Hist. Rev.: 170, pi. 5-6; Storer, 1863, Mem. Amer. Acad. 
Arts Sci. n.s. 8 (2) : 420, pi. 34, fig. 2 ; Beltremeux, 1864, Ann. Acad, la Rochelle (Faune) : 53 ; 
Couch, 1865, Hist. Fish. Brit. Is. 4: 377, pi. 245; Blanchere, 1868, Nouv. Diet, peches: 505, 
fig. 673; Schlegel, 1869, Nat. Hist. Ned. Vischen: 182, pi. 17, fig. 4; Giinther, 1870, Cat. Fish. 
Brit. Mus. 8: 317; Capello, 1870, /. Sci. Math. Phys. Nat. Lisboa 2: 136; 1881, Mem. R. Acad. 
Lisboa: 41; Andrews, 1871, Proc. Nat. Hist. Soc. Dublin (1865-1869), 5 (1) : 123; Putnam, 
1871, Proc. Amer. Ass. Adv. Sci. 19: 255; Amer. Nat. 4: 629, figs. 134, 137; Jourdain, 1871, 
C. R. Acad. Sci. Paris 63: 1225; Canestrini, 1872, Fauna d'ltalia [Pesci): 148; Barker, 1876, 
Zoologist: 5087; Malm, 1877, Goteborgs Fauna: 599, 654; Winther, 1879, Nat. Tidsskr. (3) 
12: 54; Stossich, 1879, Boll. Soc. Adriat. Sci. Nat. 5: 36; Moreau, 1881, Poiss. France 2: 74; 
Vignal, 1881, Arch. Zool. exp. gen. 9: 369, pi. 21 ; Campbell, 1883, Proc. Nat. Hist. Soc. Glasgow 
(1882) 5: 176; Day, 1884, Fish. Gt. Brit. 2: 272, pi. 148; Thompson, 1888, Anat. Anz. 3: 93, 
figs.; 1889, Stud. Mus. Zool. Univ. Coll. Dundee 1, No. 4; Vinciguerra, 1890, Boll. Mus. Zool. 
Rome 1: 33; Haller, 1891, Morph. Jb. 17: 198, figs., pis. 13-15; Steindachner, 1891, Ann. 
Naturh. Hofmus. Wien 6: 90; Almeida & Roquette, 1892, Inquir. Industr., Lisboa 2: 377; 
Girard, 1894, Ann. Sci. Nat., Porto 1:31; Tagliani, 1894, Monit. Zool. ital. 5: 248 ; Grieg, 1895, 
Bergens Mus. Aarb. 6: 11; Smitt, 1895, Skandinav. Fisk. 2: 622, figs. 153, 154 a, 156, 157, 
pi. 27, fig. 4 ; Osorio, 1896, /. Sci. Math. Phys. Nat. Lisboa 4: 157 ; Vieira, 1898, Ann. Sci. Nat., 
Porto: 24; Clarke, 1898, Zoologist 16: 439; Andersson, 1900, Ofvers. Vetensk Akad. Forh., 
Stockh.: 603; Parker, 1900, Anat. Anz. 17: 313, fig. ; Herdman & Dawson, 1902, Mem. Lanes. 
Sea Fish. Comm. 2: 57 ; Griffini, 1903, Ittiol. Ital. : 155, figs. 81, 82 ; Michailovskij , 1903, Annu. 
Mus. Zool., Acad. St. Petersb. 8:xlvi; Meek, 1904, Anat. Anz. 25: 217, fig.; Dall, 1908, Bull. 
Mus. Comp. Zool. Harv. 43 (6) : 232 ; Novikov, 1909, Dnevn. russkh. Estestroisp. 1909- 
1910: 286; 1910, Anat. Anz. 37: 97; Sauvage, 1910, Mem. Soc. Hist. Nat. Autun. 23: 1; 
Giinther, 1910, /. Mus. Godeffroy 9(17): 477; Seabra, 1911, Bull. Soc. Portug. Sci. Nat.: 193; 
Le Danois, 1913, Poiss. Manche occ: 106, fig. 182; Kaschkarov, 1916, Rev. Zool. Russe 1: no, 
figs. 1-12 ; Thompson, 1918, Scot. Nat. : 41, 59 ; Kincaid, 1919, Annot. List. Puget Sound Fish. : 
23, fig. 43 ; Toni, 1921, Atti 1st. Veneto 80: 125 ; Grenholm, 1923, Stud. Floss. Teleost. Upsala: 
240; Patroni, 1923, Ann. Mus. zool. Napoli, n.s. 5 (4), pi. 1 ; Jenkins, 1925, Fish. Brit. Is. : 212, 
pi. 85 ; Duncker & Mohr, 1926, in Grimpe & Wagler, Tierwelt Nord u. Ostsee 4 (12) : Xllg 29, 
figs. 4, 5; Gudger, 1928, Sci. Mon. N.Y.: 257; Burr, 1928, /. Comp. Neurol. 45: 33, figs.; 
Caraffa, 1929, Poiss. Corse: 50, fig.; Marine Biol. Ass. 1931, Plymouth Mar. Fauna: 318; 
Saemundsson, 1931, Nat. Reykjavik 1: 164; 1939, Vidensk. Medd. naturh. Foren. Kbk. 102: 
207; Noronha & Sarmento, 1934, Peixes Madeira: 121; Nobre, 1935, Fauna Mar. Portugal, 
Vertebr.: 240, fig. 109; Toschi, 1936, Boll. Pesca Piscicolt. Idrobiol. 12: 325 ; Sanzo, 1939, Arch. 



THE OCEAN SUNFISHES (FAMILY MOLIDAE) 115 

zool. Torino 26: 121, pi. 7, figs. 16, 17; Andersson, 1942, Fisk. Nord. 1: 62, pi.; Roon, 1942, 

Zool. Meded. 23: 313, fig. 
Orthagoriscus spinosus Cuvier, 1817, Regne Anim. ed. 2, 2: 370; Richardson, 1844, Voy. Sulphur, 

Fish.: 125, pi. 62, figs. 10-12. 
Cephalus ortagoriscus Risso, 1826, Hist. eur. Merid. 3: 173. 
Diodon carinatus Mitchill, 1828, Ann. Lyceum New York 2: 264, pi. 5, fig. 1. 
? Mola aspera Nardo, 1828, Bull. Sci. Nat. (Ferussac) 8: 437; Bonaparte, 1846, Cat. met. pesci 

eur.: 87. 
Mola hispida Nardo, 1828, ibid. : 438. 

Pedalion gigas (Guilding) Swainson, 1839, Nat. Hist. Class. Fish. 1: 199, fig. 33. 
Molacanthus pallasi Swainson, 1839, ibid. 2: 329. 

Ozodura orsini Ranzani, 1839, Novi Comment. Acad. Sci. Inst. Bonon 3: 80, pi. 6. 
Tympanomium planci Ranzani, 1839, ibid., table. 
Diplanchias nasus Ranzani, 1839, ibid. 
Trematopsis willughbii Ranzani, 1839, ibid. 

Orthragoriscus retzii Ranzani, 1839, ibid.; Bonaparte, 1846, Cat. met. pesci eur.: 87. 
Orthragoriscus ghini Ranzani, 1839, ibid. 
Orthragoriscus rondeletii Ranzani, 1839, ibid. 
Orthragoriscus blochii Ranzani, 1839, ibid. 

Orthragoriscus alexandrini Ranzani, 1839, ibid., pi. 6; Alessandrini, 1839, ibid.: 359, pis. 31-34. 
Orthragoriscus redi Ranzani, 1839, ibid., table. 
Orthragoriscus aculeatus Ranzani, 1839, ibid. 

Pallasina pallasi Nardo, 1840, Ann. Sci. Regno Lombardo-Veneto 10: 112. 
Acanthosoma carinatum De Kay, 1842, Nat. Hist. New York, Zool. 3: 330, pi. 15, fig. 179; Storer, 

1846, Mem. Amer. Acad. Arts Sci. 2: 494. 
Molacanthus hispidus Bonaparte, 1846, Cat. met. pesci eur.: 87. 
Mola luna Sassi, 1846, Saggio sopr. Pesci, Sec: 35; Aradas, 1871, Ann. Min. Agric. Ind. Comm. 

1, pt. 1 : 587. 
Orthagoriscus analis Ayres, 1859, Proc. Calif. Acad. Sci. 2: 31, fig. 14; i860, ibid.: 54, fig. 5; 

Stearns 1867, ibid. 3: 341. 
Molacanthus carinatus Gill, 1861, Proc. Acad. Nat. Sci. Philad. (i860): 21. 
Aledon storeri Castelnau, 1861, Mem. poiss. Afr. australe: 75. 
Aledon capensis Castelnau, 1861, ibid.: 76. 
Mola nasus Steenstrup & Lutken, 1863, Overs, danske Vidensk. Selsk. Fork.: 36; Wahlgren, 

1868, Acta Univ. Lund. 4: 1, pi. 
Mola retzii Steenstrup & Lutken, 1863, ibid. ; Wahlgren, 1868, ibid. 
Orthagoriscus sp. Swinhoe, 1863, Ann. Mag. Nat. Hist. (3) 12: 225. 
Orthagoriscus ozodura Harting, 1868, Verh. Akad. Wet. Amst. 11: 1, pis. 1-8. 
Orthagoriscus planci Stossich, 1879, Boll. Soc. Adriat. Sci. Nat. 5: 36. 
Orthagoriscus nasus Jeude, 1892, Notes Leyden Mus. 14: 127, pi. 5; Tijdschr. Ned. Dierk. Ver. 

18: 185, pi. 11. 
Orthagoriscus sp. Reuvens, 1894, Notes Leyden Mus. 16: 128, pi. 5. 
Mola mola Jordan, 1885, Proc. U.S. Nat. Mus. 8: 393; Eigenmann, 1893, ibid. 15 (1892): 131, 

175 ; Jordan, 1895, Proc. Calif. Acad. Sci. (2) 5: 491 ; Collett, 1896, Result. Camp. Sci. Monaco, 

10: 163 {part.); Jordan & Evermann, 1898, Bull. U.S. Nat. Mus., No. 47, 2: 1753; H. M. 

Smith, 1898, Bull. U.S. Fish. Comm. 17: 85; Linton, 1898, Proc. U.S. Nat. Mus. 20: 507 et 

seq. ; Evermann & Kendall, 1899, Rep. U.S. Fish. Comm.: 88; Jordan & Snyder, 1901, Proc. 

U.S. Nat. Mus. 24: 260; Green, 1901, Bull. U.S. Fish. Comm. 19*. 321 ; Jordan & Evermann, 

1902, Amer. Food and Game Fish.: 492, fig.; Gilbert & Starks, 1904, Mem. Calif. Acad. Sci. 

4: 206; Hargitt, 1905, Bull. U.S. Bur. Fish. 24 (1904): 25; Stead, 1906, Fish. Austral.: 227, 

fig. 82; Starks & Morris, 1907, Univ. Calif. Publ. Zool. 3 (n): 205; Murray & Hjort, 1912, 

Depths of the Ocean: 644; Halkett, 1913, Checklist Fish. Canada: 116; Dean, 1913, Amer. Mus. 

J. 13 (8) : 370, fig. ; Hilton, 1914, J . Ent. Zool. 6(4): 233; Evermann, 1915, Copeia, 20: 17; 

Buen, 1919, Bol. Pesc. Madr. 4: 295 ; 1935, Notas. Inst. esp. Oceanogr. 2 (89) : 146; Dons, 1920, 



n6 THE OCEAN SUNFISHES (FAMILY MOLIDAE) 

Troms. Mus. Adrsh. 43 (6): 38, pi. 2; Jordan, 1921, Copeia, 93: 28; McCulloch, 1922, 
Aust. Zool. 2: 130, pi. 43, fig. 374 a; Fowler, 1923, Proc. Acad. Nat. Sci. Philad. 75: 294; 
Wolleboek, 1924, Norges Fiske: 224, fig. 254; Damant, 1925, Nature, Lond. 116: 543, rig.; 
Bigelow & Welsh, 1925, Bull. U.S. Bur. Fish. 40 (1) : 301 ; Buen, 1926, Result. Camp. int. Inst, 
esp. Oceanogr. 2: 56; Barnard, 1927, Ann. S. Afr. Mus. 21: 986; Fowler, 1928, Mem. Bishop 
Mus. 10: 473; Ulrey & Greeley, 1928, Bull. Calif. Acad. Sci. 27 (1) : 24; Breder, 1929, Field 
Book Mar. Fish. Atlant. Coast: 236, fig.; Hubbs & Schultz, 1929, Calif. Fish Game, 15 (3): 
Ulrey, 1929, /. Pan-Pacif. Res. Inst. 4 (4) : n, 235 ; McCulloch, 1930, Mem. Aust. Mus. 5: 436; 
Myers & Wales, 1930, Copeia 1934: 11 ; Ancona, 1931, Faune Flore Mediter., figs. 1, 2 ; Breder, 
1932, Copeia (4) : 180 ; Gregory, 1933, Tnms. A mer. Phil. Soc. 23 (2) : 294 ; Gregory & Raven, 1934, 
Copeia^: 145; Barnard, 1935, Ann. S. Afr. Mus. 30: 645; Barnhart, 1936, Mar. Fish. South. 
Calif. : 95, fig. 288 ; Tibby, 1936, Calif. Fish Game 22 (1) : 49, fig. 22 ; Fowler, 1936, Bull. Amer. 
Mus. Nat. Hist. 170 (2) : 1123, fig. 469; Schultz & De Lacy, 1936, Mid-Pac. Mag. 49 (3) : 211 ; 
Scofield, 1937, Calif. Fish Game 23 (4): 336; Schultz, 1938, Nat. Geogr. Mag. 74 (4): 497; 
Brimley, 1939, /. Elisha Mitchell Sci. Soc. 15 (2) : 301, pi. 30; Deranyigala, 1944, /. Bombay 
Nat. Hist. Soc. 44 (3) : 429 ; Mendes, 1944, Bol. Fac. Filos. Cien. Let. Univ. S. Paula, Zool. No. 
8: 173, pi. ; Engel, 1945, Zool. Meded. Leiden 25: 11, pi. 1 ; Clemens & Wilby, 1946, Bull. Fish. 
Res. B. Canada 68: 330, fig.. 247; Medcof & SchifTman, 1947, Acadian Nat. New Brunswick 2: 
8, 63, fig.; Poll, 1947, Poiss. Mar.: 405, figs. 260, 261; Barnard, 1948, Ann. S. Afr. Mus. 
36 (5): 401, pis. 12, 13; Maul, 1949, Vertebr. Madeira, ed. 2, 2 (Peixes) : 158; Clark, 1949, 
Amer. Mus. Novit. 1397: 7, fig. 9; J. L. B. Smith, 1949, Sea Fish. S. Africa: 422, pi. 95, 
fig. 1213; Tortonese, 1950, Att. Ace. Ligure Sci. 6 (1): 112. 

Orthragoriscus nasus Reuvens, 1897, Notes Ley den Mus. 18: 209, pi. 3. 

Mola ramsayi Whitley, 1931, Rec. Aust. Mus. 18 (3) : 126 (part.), fig. 2, pi. 16, fig. 1 ; 1933, Vict. 
Nat. 49: 210, figs. 1, 2 (not of Giglioli). 

Mola alexandrini Barnard, 1948, Ann. S. Afr. Mus. 36 (5): 402. 

The above extensive synonymy illustrates the considerable literature which has 
accumulated concerning this species. From a perusal of this data it is possible to give 
a rather more complete account than for other members of the family, but there is 
still much of its biology that remains conjectural. The anatomy has been studied 
broadly and in detail by a number of workers, and from this, together with descrip- 
tions or figures giving reliable information about the clavus, it seems quite clear that 
not more than one species is involved. Published records, considered statistically, 
would give the impression that the species is mainly a North Atlantic one, becoming 
rarer southwards, in the Indian Ocean and in the Western Pacific, but this is possibly 
an illusion due to the much higher rate of publication in the Atlantic and Mediter- 
ranean countries. 

Certainly the Japanese form is not separable from the Atlantic form, since we have 
specimens from Japan in our collection for comparison; according to Jordan and 
Fowler it occurs at Hawaii, and it seems to be common at California, so that it is 
replaced by M. ramsayi only in the South Pacific. I am much indebted to Mr. W. I. 
Follett, of the California Academy of Sciences, for information and radiographs which 
enable me to identify the Calif orni an specimens. 

A bad practice among some authors is the borrowing of an illustration from some 
earlier work, especially when the specimen depicted was obtained in a locality remote 
from that being discussed. Mola mola has suffered much from this treatment, and 
in consequence it is not possible to be definite as to the identity of specimens in 
regions where M. ramsayi might occur also, because the distinguishing characters 



THE OCEAN SUNFISHES (FAMILY MOLIDAE) 



117 



of the clavus have been hitherto unknown and are, therefore, not described ; a re- 
liable picture might have given the answer. 

Comparison of adequate descriptions and figures shows that some order underlies 
the variability which has been remarked upon by so many authors. After meta- 
morphosis the young fishes are short and deep, the snout not protuberant, the fins 
rather narrow, and the margin of the clavus is not conspicuously lobed. The length 
of the clavus from the posterior edge of the 'carapace' — i.e. the anterior edge of the 
smooth band between dorsal and anal fins — is much less than that of the head. When 
the fish exceeds a length of about 2 ft., however, sexual differences become apparent. 
The bony tubercle on the snout is either pushed forward (in the male) , or upward (in 
the female) ; in consequence the male develops a 
pronounced snout, projecting forward (the 'nasus' 
form), while the female appears more deep-headed, 
with the front of the snout nearly vertical (the 
4 alexandrint form). As growth proceeds the clavus 
develops backwards between the ossicles, forming a 
series of lobes which at first number between 9 and 
12 in both sexes ; females do not seem to pass beyond 
this stage, but in large males the five median lobes 
become very large and the others reduced. After 
the formation of the lobes the clavus is probably 
always longer in a male than in a female of the same 
size, and in the biggest males it may be as long as the 
head. In large specimens of both sexes two prominent, 
swollen ridges are formed on each side of the head ; 
these are discernible in small examples, and are 
evidently analogous if not homologous with the late- 
ral ridges of Ostracionts, but with age they become 
very conspicuous. In the larger examples also the 
dorsal and anal fins are relatively much broader. 

All this is indicated by a study of the records. 
Comparatively few of the specimens described have 

been examined for sex, but in each case where the sex is stated the characters 
mentioned above are found to be associated with it ; of particular interest is the 
paper by Roon & Pelkwijk (1939), who had both sexes and figured them together. 
Harting's (1868) plate 1 gives a fair representation of a female, and Whitley 
(1931) has given a drawing of another, together with a photograph of it (pi. xvi, fig. 1), 
which shows the lateral ridges excellently, and Murray & Hjort's (1912) photograph, 
copied by Schmidt, illustrates a fine male. The various phases of development out- 
lined do not always coincide with a particular size or age, but are evidently dependent 
to some extent on environmental circumstances. 

Mola mola grows to a great size, the largest record being apparently that by Dean 
(1913), measuring 10 ft. 1 in. in length and n ft. from tip of dorsal fin to tip of anal 
fin, a male. Mikailovskij (1903) described one measuring 8 ft. 6 in. in length and 
weighing 1,410 kg. Jeude (1890) described a specimen 2-23 m. (7 ft.) in length, 




Fig. 15. Post-larvae of Mola. A. 
' Ostracion hoops' stage (5 mm.). 
(After Schmidt) ; B. ' Molacanthus' 
stage (16 mm.). (From specimen 
in the British Museum collection.) 



n8 



THE OCEAN SUNFISHES (FAMILY MOLIDAE) 



apparently a female. The specimen recorded by Gunther as ' 7 feet long, Portsmouth ' 
was the fish taken by Parlby (1849), who described its capture at Chesil Beach and 




Fig. 16. Mola mola, adult, 600 mm. long, Plymouth. (From stuffed specimen 

in the British Museum collection.) 

na, area of reduced denticles; po, paraxial ossicle. 

stated that it measured 6 ft, 3 in. long. It was probably a male. As a stuffed skin it 
remained in the British Museum collection until recently, when it was found to be in 
a bad state and destroyed ; my (calliper) measurement at this time reading 5 ft. 8 in., 
the loss being presumably due to shrinkage (unless Parlby made a contour measure- 



THE OCEAN SUNFISHES (FAMILY MOLIDAE) 



119 



ment) . A number of smaller stuffed skins and several specimens in spirits remain in 
the collection. It is never common, the large literature being due to the great interest 




Fig. 17. Mola mola, young adult, 366 mm. long, Chouse, Japan. (From 
specimen in spirits in the British Museum collection.) 

it arouses, almost every specimen being reported upon ; but it is more frequently met 
with than any of the previous species. Nevertheless its early developmental stages 
are less well known than those of Ranzania, and fertile eggs or early larvae have not 
been found ; it is not improbable that it spends a great part of its life in deep water. 



120 THE OCEAN SUNFISHES (FAMILY MOLIDAE) 

The scarcity of young specimens is remarkable when we consider that a female 
4 ft. 6 in. long contained 300 million eggs. The mode and place of breeding have yet 
to be found. 

Its migrations inshore are unpredictable, and are usually supposed to coincide with 
invasions of medusae, salps, and ctenophores, upon which it largely feeds. Specimens 





Fig. 18. Different lobulation of the clavus, with similar skeletal 

supports, in Mola mola. Drawn to the same size for comparison. 

That on the right is the characteristic form in large males. 

taken inshore, however, are usually found to be feeding on littoral forms, and the list 
of organisms taken from stomachs includes Crustacea, ophiuroids, molluscs, hydroids, 
ctenophores, corallines, and algae ; Schmidt has reported them as feeding heavily on 
leptocephali ; on one occasion a flounder [Platichthys flesus) was found in the throat 
(Reuvens, 1897), and in our collection there is a ling (Molva macrophthalma) two feet 
long which was taken from the stomach of Mola mola. The stomach is not infrequently 
found to be empty, and it is quite probable that the specimens so frequently taken 
without difficulty while 'basking' at the surface are in fact sick or dying fish. Myers 
& Wales (1930) found young fish to be active and alert, but later found two larger fish 
'disabled' at the surface. It would be interesting to know the cause of such disable- 
ment . Possibly the great variety of parasites with which they are often found to be 
infested may have some bearing on the matter. 



THE OCEAN SUNFISHES (FAMILY MOLIDAE) 121 

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23-25. (Molidae, p. 635, figs. 5-7.) 

Chabanaud, P. 1935. Quelques Monstruosites chez des Poissons HeteYosomes. Arch. Mus. 

Hist. Nat. Lyon, 15: 1-23, 4 pis. 
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10: 1-18. 
Gregory, W. K., & Raven, H. C. 1934. Notes on the Anatomy and Relationships of the Ocean 

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Bull. U.S. Fish Comm. 19: 321. 
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1937 °- The Natural History and Geographical Distribution of the Pointed-tailed Sunfish 

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Opisthoproctus soleatus Vaillant. Proc. Zool. Soc. Lond. (3) : 601-614, 2 pis. 
Whitley, G. P. 1931. Studies in Ichthyology No. 4. Rec. Aust. Mus. 18 (3) : 96-133, pis. 11-16. 

(Molidae, p. 126, fig. 2, pi. 16.) 




PRESENTED 

3 1 OCT 1951 






PRINTED IN 

GREAT BRITAIN 

AT THE 

UNIVERSITY PRESS 

OXFORD 

BY 

CHARLES BATEY 

PRINTER 

TO THE 
UNIVERSITY 



2 JUL 1952 

HE CESTODES OF 

SEALS FROM 
THE ANTARCTIC 



STANISL-AW MARKOWSKI 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ZOOLOGY Vol. i No. 7 

LONDON : 1952 



THE CESTODES OF SEALS FROM 
THE ANTARCTIC 

BY 

STANISLAW MARKOWSKI 

Associate, Department of Zoology 
British Museum (Natural History) 




Pp. 123-150; Pis. 10-21 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ZOOLOGY Vol. 1 No. 7 

LONDON : 1952 



THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted in 1949, is issued 
in five series, corresponding to the Departments of the 
Museum. 

Parts appear at irregular intervals as they become ready. 
Volumes will contain about three or four hundred pages, 
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year. 

This paper is Vol. 1, No. 7, of the Zoological series. 



PRINTED BY ORDER OF THE TRUSTEES OF 
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Issued January 1952 Price Twelve shillings and sixpence 



THE CESTODES OF SEALS FROM THE 

ANTARCTIC 

By STANISLAW MARKOWSKI 

ASSOCIATE, DEPARTMENT OF ZOOLOGY, BRITISH MUSEUM (NATURAL HISTORY) 

SYNOPSIS 

The present paper reviews and re-describes all known Pseudophyllidean Cestodes occurring in the five 
species of Antarctic Seals, namely, Weddell seal, Leopard seal, Crabeater seal, Elephant seal, and Ross 
seal. According to the literature, twelve species belonging to two genera have been previously recorded 
from these hosts. 

The investigation of material collected by the British Graham Land Expedition, together with a 
comparative study of type-specimens collected by other Antarctic expeditions, leads to the conclusion that 
there are nine species of Pseudophyllidean Cestodes occurring in these hosts. These belong to four genera, 
two of which are new and of these, one represents a new species. 

From the twelve species quoted in the literature, three are here placed in synonymy and one transferred 
to another genus. 

MATERIAL AND METHODS 

The bulk of the material for this study was collected by the British Graham Land 
Expedition during 1934-1937. In addition, some samples gathered by the Discovery 
in 1925, 1928, and 1931, and by the 'Falkland Islands Dependencies Survey' in 1945, 
have been examined for comparative purposes. The material obtained by the British 
Graham Land Expedition comprises 33 lots from Weddell seals (Leptonychotes 
weddelli), 12 from Leopard seals (Hydrurga leptonyx), 6 from Crabeater seals (Lobodon 
carcinophagus) , and 2 from Elephant seals (Macrorhinus leoninus). The Discovery 
material consists of 5 lots from Leopard seals and 1 from a Crabeater seal, and that of 
the ' Falkland Islands Dependencies Survey* of 2 lots only from Weddell seals. Thus 
the 61 batches of specimens investigated can be summarized as follows: 35 from 
Weddell seals, 17 from Leopard seals, 7 from Crabeater seals, and 2 from Elephant 
seals. 

The British Graham Land Expedition's material was obtained from Graham Land 
(Debenham Is., Horseshoe I., Argentine Is., and Beascochea Bay), South Shetland 
(Deception I.), Palmer Archipelago (Melchoir I.), and South Georgia (Cooper Bay 
and Bay of Isles). The Discovery material was collected from South Orkneys 
(Coronation I.), South Georgia (Maivicken), and South Sandwich Is. The 'Falkland 
Islands Dependencies Survey' from Graham Land (Hut Cove, Hope Bay) and 
Palmer Archipelago (Port Lockroy, Wiencke I.). Preservation was in 4 per cent, 
formalin or, occasionally, in Bouin's Solution. The material comprises mainly 
portions of the gut with Cestodes attached to the walls, and individual specimens are, 
with a few exceptions, perfectly extended. 

In addition to the new material, numerous collections from the same host-species 
were examined. These included the type-specimens described by Baird (1853), 
Shipley (1907), Rennie & Reid (1912), and Leiper & Atkinson (1914). The collection 



126 THE CESTODES OF SEALS FROM THE ANTARCTIC 

of Fuhrmann consisted of microscopical preparations of specimens, probably types, 
of Linstow (1892), of Railliet & Henry (1912), and material described by Fuhrmann 
(1920) himself. Some of the material in these collections is not in very good condi- 
tion, the specimens mounted whole or as serial sections having partially lost their 
stains. 

Over 1,200 slides of the present material have been made. Whole preparations 
have been stained with Mayer's paracarmine or alum carmine, and serial sections, 
10 fM to 18 ju, in thickness, have been double-stained with Ehrlich's haematoxylin and 
erythrosin. 

I have great pleasure in expressing my heartiest thanks to Dr. H. A. Baylis, who 
kindly suggested this investigation and provided the necessary materials ; to Dr. 
H. W. Parker, Keeper of the Department of Zoology, British Museum (Natural 
History) ; to Dr. M. Burton; and to Mr. S. Prudhoe for his valuable assistance. In 
addition, I wish to express my thanks to Professor J. G. Baer, Rector of the University 
of Neuchatel, for his kindness in lending me the collection of slides left by the late 
Professor O. Fuhrmann, and to Dr. G. C. L. Bertram, of St. John's College, Cam- 
bridge, for information on the Graham Land Seals. 

HISTORICAL 

Our knowledge of Cestodes occurring in the Antarctic seals has been obtained 
almost exclusively from various Antarctic expeditions, including five British, one 
Australian, one French, and two German expeditions. 

The first specimens were collected in 1 839-1 843 by Ross's Antarctic Expedition 
from a Phoca sp. — probably the Ross seal — and described by Baird (1853) as Bothrio- 
cephalus antarcticus. Further samples were collected from the Ross seal by the 
National Antarctic Expedition [Discovery) in 1901-1904 and described by Shipley 
(1907) under the name Dibothriocephalus antarcticus. Railliet & Henry (1912) studied 
this species from material collected from the Ross seal by the Second French Antarctic 
Expedition of Dr. J. Charcot (Pourquoi Pas ?) 1908-1910 and described it as Diphyllo- 
bothrium antarcticum. Finally, Fuhrmann (1920) l re-examined the type-specimens 
and made the species the type of a new genus, Glandicephalns , characterized by the 
peculiar development of the musculature of the body, by the arrangement of the 
testes, and by the presence of gland-cells in the scolex. Johnston (1937) mentions 
the species as having been found in the Ross seal by the Australasian Expedition, 
1911-1914. 

Linstow (1892) described as Bothriocephahis tectus headless specimens from 
Elephant seals, collected at South Georgia by the German Expedition in 1882-1883. 
This was also recorded by Linstow in Shipley (1902) from the Ross seal, collected by 
the Southern Cross Expedition, 1898-1900, and by Johnston (1937) from the Elephant 
seal, in the collection of the Australasian Antarctic Expedition. 

Diphyllobothrium quadratum (Linstow 1892) was collected for the first time from 
the Leopard seal by the German Expedition, 1882-1883, in South Georgia and in 
1902-1904 by the Scottish National Antarctic Expedition [Scotia). It was re-described 

1 Fuhrmann's paper was published in 1920 (December) and not in 1921, as is often quoted. 



THE CESTODES OF SEALS FROM THE ANTARCTIC 127 

by Rennie & Reid (19 12) as Bothriocephahts coatsi, from the same host. Railliet & 
Henry (191 2) recorded this species as Diphyllobothrium resimum, collected by the 
Second French Antarctic Expedition. Later, Fuhrmann (1920) gave a description of 
D. quadratum, collected by the German South Pole Expedition, 1901-1903, and 
regarded Bothriocephalus coatsi (Rennie & Reid, 1912) and Diphyllobothrium resimum 
(Railliet & Henry, 1912) as synonyms of that name. Finally, Johnston (1937) gave 
a few further details of the species, based on material from the Leopard seal, collected 
by the Australasian Antarctic Expedition. 

Shipley (1907) described Diphyllobothrium wilsoni and D. scotti, the first being 
obtained from the Weddell seal and the Ross seal, and the second from the Ross seal, 
both having been collected by the National Antarctic Expedition, 1901-1904. They 
were re-described by Fuhrmann (1920) from the material obtained by the German 
South Pole Expedition (1901-1905) from the Ross seal and Leopard seal. They are 
also reported by Johnston (1937) from the Ross seal and the Weddell seal collected 
by the Australasian Antarctic Expedition. 

Diphyllobothrium mobile (Rennie & Reid, 1912) from the Weddell seal and D. 
scoticum (Rennie & Reid, 1912) from the Leopard seal were collected for the first 
time by the Scottish National Antarctic Expedition in 1902-1904 and described by 
Rennie & Reid (1912) under the generic name Dibothriocephalus. Fuhrmann (1920) 
gives a full re-description of both these species from the Ross seal and the Weddell 
seal, and of D. scoticum from the Leopard seal, collected by the German South Pole 
Expedition in 1901-1903. Both species were studied by Johnston (1937) from material 
from the Weddell seal and the Leopard seal, collected by the Australasian Antarctic 
Expedition. 

Diphyllobothrium per joliatum Railliet & Henry, 1912, described also as D. clavatum 
in the same work, was collected for the first time by the Second French Antarctic 
Expedition from the Weddell seal. It was re-described by Leiper & Atkinson (1915) 
from material collected by the British Antarctic [Terra Nova) Expedition 1910 and 
by Fuhrmann (1920) from material collected by the German South Pole Expedition 
under the name Dibothriocephalus perfoliatus. In both cases the material was taken 
from the same host. Fuhrmann (1920) recognized D. clavatum as a synonym of D. 
perfoliatum. This Cestode is also mentioned by Johnston (1937) in a collection made 
by the Australasian Antarctic Expedition. 

Diphyllobothrium rufum Leiper & Atkinson, 19 14, collected by the Terra Nova 
Expedition in 1910, is considered by Johnston (1937) as 'a short-necked, precocious 
form of D. perfoliatum', found in the Weddel seal by the Australasian Antarctic 
Expedition. 

The last two species, Diphyllobothrium lashleyi and D. archeri, were both described 
by Leiper & Atkinson (19 14) from the Weddell seal gathered by the Terra Nova 
Expedition. The first of these species was re-described by Johnston (1937) from 
material from the same host-species collected by the Australasian Antarctic 
Expedition 1911-1914. 

To sum up, 9 species have been described from the material collected by the British 
Antarctic Expeditions, 4 by the French, and 2 by the two German Expeditions. 

The details of the result of each expedition are shown in Table No. 1. 



128 



THE CESTODES OF SEALS FROM THE ANTARCTIC 



Table No. i 

A list of the Cestodes from Seals, collected by Antarctic 
Expeditions during the period i8jg-igi4 



Expeditions 


Parasite 


Host 


Ross's Antarctic Exp. 


Glandicephalus antarcticus (Baird, 1853) 


Phoca sp., probably 


1839-1843 






Ross seal 


Southern Cross Antarctic 


Diphyllobothrium tectum (Linstow, 1892) 


Ommatophoca rossi 


Exp. 1 898-1900 








National Antarctic Exp. 


,.. 


scotti (Shipley, 1907) 


,, 


(Discovery) 1 901-1909 


,, 


wilsoni (Shipley, 1907) 


,, 




Glandicephalus 


antarcticus (Baird, 1853) 


,, 


Scottish National Antarctic 


,, 


antarcticus (Baird, 1853) 


,, 


Exp. (Scotia) 1902-1904 


Diphyllobothrium coatsi (Rennie & Reid, 1912) 


Hydrurga leptonyx 




., 


mobile (Rennie & Reid, 191 2) 


Leptonychotes weddelli 




•' 


scoticum (Rennie & Reid, 


Hydrurga lepionyx 




Phyllobothrium 


1912) 
sp. (larva) Rennie & Reid, 191 2 


Leptonychotes weddelli 


British Antarctic Exp. 


Diphyllobothrium mobile (Rennie & Reid, 191 2) 


,, 


(Terra Nova) 1910-1913 


,, 


coatsi (Rennie & Reid, 1912) 


,, 




» 


perfoliatum (Railliet & Henry, 


" 




» 


1912) 
archeri (Leiper & Atkinson, 
1914) 


" 




>» 


lashleyi (Leiper & Atkinson, 
1914) 


" 




" 


rufum (Leiper & Atkinson, 
1914) 


" 


Australasian Antarctic 


,, 


perfoliatum Railliet & Henry, 


,, 


Exp. 1911-1914 


,, 


1912 
lashleyi (Leiper & Atkinson, 

1914) 
wilsoni (Shipley, 1907) 


,, 




tt 


„ 




>> 


rufum Leiper & Atkinson, 


•• 






1914 
mobile (Rennie & Reid, 191 2) 






,, 


quadratum (Linstow, 1892) 


Hydrurga leptonyx 




,, 


scoticum (Rennie & Reid, 


,, 






1912) 






„ 


scotti (Shipley, 1907) 


Ommatophoca rossi 




>> 


wilsoni (Shipley, 1907) 
mobile (Rennie & Reid, 1912) 


" 




Glandicephalus antarcticus (Baird, 1853) 


,, 




Diphyllobothrium tectum- (Linstow, 1892) 


Mirounga leonina 




Phyllobothrium 


sp. (larva) 


.. 


German Antarctic Exp., 


Diphyllobothrium tectum (Linstow, 1892) 


Mirounga leonina 


South Georgia, 1882-1883 


,, 


quadratum (Linstow, 1892) 


Hydrurga leptonyx 


German Antarctic Exp. 


,, 


perfoliatum Railliet & Henry, 


Leptonychotes weddelli 


(Gauss) 1901-1903 




1912 






,, 


quadratum (Linstow, 1892) 


Hydrurga leptonyx 




" 


wilsoni (Shipley, 1907) 


Ommatophoca rossi, 
Hydrurga leptonyx 




" 


mobile (Rennie & Reid, 191 2) 


Ommatophoca rossi, 
Leptonychotes weddelli 



THE CESTODES OF SEALS FROM THE ANTARCTIC 

Table No. i (continued) 



129 



Expeditions 


Parasite 


Host 


2nd French Antarctic Exp. 
{Pourquoi Pas?) 1908- 
191 0, Dr. J. Charcot 


Diphyllobothrium resimum Railliet & Henry, 
1912 
,, wilsoni (Shipley, 1907) 
,, perfoliatum Railliet & Henry, 


Hydrurga leptonyx 
Leptonychotes weddelli 




i» 


clavatum Railliet & Henry, 


» 




sp. ? Railliet & Henry, 1912 
Glandicephalus antarcticus (Baird, 1853) 
Cestoda (unidentified) 


Ommatophoca rossi 
Lobodon carcinophagus 



The species enumerated above have been listed by Meggitt (1924) and Stunkard 
& Schoenborn (1936), though all these authors appear to have overlooked the work 
of Fuhrmann (1920), who reduced the number of species to twelve, apportioned 
between two genera, Diphyllobothrium and Glandicephalus. 1 

Wardle, McLeod, & Stewart (1947) have proposed a new classification of the genus 
Diphyllobothrium, but this appears to have been based mainly upon information 
obtained from the literature. Stunkard's (1948) criticism of the classification is fully 
subscribed to by the present writer. 

DISCUSSION 

After investigation of the new material it seems that the Pseudophyllidean Cestodes 
occurring in the Antarctic seals represent no more than 4 genera and 9 species 
namely : 

1. Diphyllobothrium lashleyi (Leiper & Atkinson, 1914). 

2. D. mobile (Rennie & Reid, 1912). 

3. D. quadratum (Linstow, 1892). 

4. D. scoticum (Rennie & Reid, 1912). 

5. D. wilsoni (Shipley, 1907). 

6. Glandicephalus antarcticus (Baird, 1853). 

7. G. [Diphyllobothrium] perfoliatus (Railliet & Henry, 1912). 

8. Baylisia baylisi gen. nov., spec. nov. 

9. Baylisiella tecta (Linstow, 1892) gen. nov. 

The details of their anatomical differences are given in Table No. 2. 

Of the five species of Diphyllobothrium mentioned above, D. lashleyi alone possesses 
a well-developed distinct neck and the rudiments of genital organs at some distance 
behind it. In this it resembles both species of Glandicephalus. The rest of the species 
of Diphyllobothrium possess a very short indistinct neck, the presence of which has 

1 In addition, unidentified Cestodes have been recorded from the Crabeater seal by Railliet & Henry 
(1912) ; and the larval stages of Phyllobothrium in the blubber of the Weddell seal by Rennie & Reid 
(1912) and Fuhrmann (1931). Phyllobothrium delphini (Bosc, 1802) Gervais, 1885, found by J. E. Hamilton 
in 1931 in the blubber of the Leopard seal at Falkland Islands is reported by Southwell & Walker (1936), 
and the larval stage of Phyllobothrium from Elephant seal by Johnston (1937). 



13° 



THE CESTODES OF SEALS FROM THE ANTARCTIC 



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PLATE 10, FIGS. 1-9 

Abbreviations used: c, cirrus; c.b., 'calcareous body'; e.g., cephalic glands; 
c.s., cirrus-sac; d.v.m., dorso-ventral musculature; ex, excretory system; g, glands; 
g.p., genital papillae; l.m., longitudinal musculature; m, muscles; n, nerve; o, ovary; 
oc., oocapt; r.s., receptaculum seminis; s.g., shell-gland; t., testis; t.m., transverse 
musculature; u., uterus; v., vagina; v.d., vas deferens; v.g., vitelhne glands; v.s., 
vesicula seminalis; y.r., yolk reservoir. 

Fig. i. Diphyllobothrium lashleyi from Weddell seal. Sagittal section 
of cirrus-sac. 

Fig. 2. D. mobile from Weddell seal. Sagittal section of cirrus-sac. 

Fig. 3. D. quadratum from Leopard seal. Sagittal section of cirrus-sac. 

Fig. 4. D. scoticum from Leopard seal. Sagittal section showing 
glandular cells. 

Fig. 5. D. wilsoni from Weddell seal. Sagittal section of cirrus-sac, 
with coiled cirrus showing muscle-fibres attached to the dorsal wall 
of the segment. 

Fig. 6. Glandicephalus antarcticus from Ross seal. Sagittal section of 
cirrus-sac. 

Fig. 7. Glandicephalus perfoliatus from Weddell seal. Sagittal section 
of cirrus-sac. 

Fig. 8. Baylisia baylisi from Crabeater seal, (a) Sagittal section of 
cirrus-sac. (b) Vesicula seminalis in the same section. 

Fig. 9. Baylisiella tecta from Elephant seal. Sagittal section of cirrus- 



PLATE 10 




zoo. i. 7. 



Figs. 1-9 
Q 



THE CESTODES OF SEALS FROM THE ANTARCTIC 131 

often been questioned by previous authors. The neck of these species of Diphyllo- 
bothrium is extremely short, no more than a continuation of the scolex, and the rudi- 
ments of the genital organs occur in the immediate vicinity of the neck, usually in 
the second segment. 

Diphyllobothrium perfoliatum Railliet & Henry, 1912, has been transferred by the 
present writer to the genus Glandicephalus Fuhrmann (1920), owing to the peculiar 
structure of its longitudinal musculature, which is almost identical with the muscu- 
lature in G. antarcticus. In addition, there is a similar, though not identical distribu- 
tion of the testes. Both these features were stressed by Fuhrmann (1920) in his 
original diagnosis. Both species also possess imbricated proglottids, though these 
are less developed in G. antarcticus. The name Glandicephalus is not very appropriate, 
as cephalic glands may also occur in species of Diphyllobothrium. On the other hand, 
cephalic glands have not been found in the scolex of Glandicephalus perfoliatus 
stained with Ehrlich's haematoxylin. Nevertheless, on anatomical grounds it finds 
its closest affinities in G. antarcticus. 

It would seem that the cirrus-sac of the male genital apparatus, examined in 
sagittal section, and the longitudinal musculature, seen in transverse section, give 
the best means of determining species. This is true not only for Diphyllobothrium 
but for the other forms dealt with here. Earlier writers paid much more attention to 
the size of the body, shape and size of the scolex and of the neck, and the number, 
shape, and size of the segments. 

More attention was paid to the female genital system than to that of the male. 
Such features as the shape and number of the uterine coils, the size and shape of the 
ovary, and the size of the eggs and vitelline glands were considered significant, as 
well as the position of the external openings of the genital apparatus. In some cases 
the presence of cephalic glands in the scolex was accepted as a generic difference 
(Nybelin, 1931). 

There is nothing more deceptive in the species of Diphyllobothrium than the size of 
the body, its length and width. For example, D. lashleyi may vary from 4-5 cm. to 
19-5 cm. in length in the adult. Greater differences in length, observed in new material, 
have been recorded for D. scoticum, which varies from 13 cm. to 130 cm., and for 
D. wilsoni from 10 mm. to 50 mm. The number of segments also is variable. 

The scolex is also of a doubtful value, as its size and shape depends much on the 
fixatives used. Only in a few cases does it present distinct morphological differences. 

The presence of cephalic glands in the scolex has been demonstrated in D. lashleyi, 
D. mobile, and D. quadratum by using Ehrlich's haematoxylin. They were also 
found to occur in the scolex of Glandicephalus antarcticus by Fuhrmann (1920) 
and by Nybelin (1931) in Adenocephalus. This means that they occur in widely 
different genera. This feature cannot therefore be used for generic distinction. Only 
in D. scoticum, Baylisia baylisi, and Baylisiella tecta does the scolex show character- 
istic differences as compared with the other species mentioned in this paper. 

The neck is extremely variable in size, and possesses a diagnostic value only when 
it constitutes a well-marked feature. As previously stated, a neck is present in D. 
lasheyi and is also fairly well developed in Glandicephalus antarcticus and G. perfoliatus. 
In D. lashleyi, however, it is so contracted in some cases that the specimen gives an 



i 3 2 THE CESTODES OF SEALS FROM THE ANTARCTIC 

impression of being without a neck and, as a consequence, of belonging to another 
species. 

Owing to this variation of the neck Leiper & Atkinson (1914) described Diphyllo- 
bothrium rufum as a separate species, but it now appears to be a synonym of G. 
perfoliatus. D. rufum was distinguished by the absence of a neck, an effect which 
might be caused by contraction, and by ' notches ' in the imbricated portions of the 
segments. 

The female genital apparatus is of little use for purposes of identification. The 
uterus is a more or less irregular sac, rilled with eggs, with the opening, in the 
majority of cases, irregularly alternate. Only in D. scoticum is the terminal part of 
the uterus modified to form a characteristic ' pocket ' lined with villous tissues. 

The ovary also varies considerably in shape and size, even in the same strobila. 
In D. lashleyi, for instance, the ovaries in squarish segments are entirely different in 
shape to those in elongate segments. 

The sizes of eggs, given in Table No. 2, do not show any remarkable difference, 
except in D. scoticum, but this differs from other species in many other anatomical 
details, much more distinct than the size of the eggs. 

The vitelline glands, forming a continuous field along the strobila and covering 
internal structures in all species, have a characteristic form only in D. mobile, where 
this continuous field is interrupted by the transverse segmentation of the body. 

The excretory system, proposed as a criterion for identification of some of the 
Cestodes from seals by Zschokke (1903), is also uncertain since it undergoes changes 
during the fixation and the consequent contraction of the body. 

An excretory system occurs in the cortical and medullary parenchyma in the 
Cestodes of the Antarctic seals belonging to the species of Diphyllobothrium, although 
in the present material of D. mobile no excretory system has been detected. In 
Baylisia baylisi and Baylisiella tecta the system seems to be present only in the 
cortical parenchyma, where it reaches a high degree of development. 

The most essential specific differences occur in the male genital apparatus and in 
the development of the longitudinal musculature, as well as in the relation of this 
musculature to the transverse and the dorso-ventral muscles. The cirrus-sac, 
examined in sagittal section, differs specifically in shape and size. Plate 10, figs. 
1-9, shows these morphological differences exhibited by the various species enumer- 
ated in this paper. The size, shape, and position of the vesicula seminalis in relation 
to the cirrus-sac may be very variable in some species, but in others it is sufficiently 
constant to have a taxonomic value. The testes, which in the species of Diphyllo- 
bothrium occur in two lateral fields, are arranged in a single layer and are usually 
confluent in the anterior part of the segment. In D. mobile, however, they are arranged 
in one field in the anterior part of the segment, and in D. scoticum they form two 
separate fields, one at each side of the segment. The two species of Glandicephalus 
have testes scattered more or less irregularly. The same arrangement occurs in 
Baylisiella tecta. In the last case the precise number of testes occurring in the 
segment is difficult to determine, the more so as they are arranged in several irregular 
layers. The number of testes per segment has not, therefore, any value as a taxonomic 
criterion in Baylisiella. The numbers of the testes counted in transverse and sagittal 



PLATE 11, FIGS. 10-18 

(For list of abbreviations see Plate 10) 

Fig. io. Diphyllobothrium lashleyi f rom Weddell seal. Transverse section 
of muscular system. 

Fig. ii. D. mobile from Weddell seal. Transverse section of muscular 
system. 

Fig. 12. D. quadratum from Leopard seal. Transverse section of muscu- 
lar system. 

Fig. 13. D. scoticum from Leopard seal. Transverse section of muscular 
system. 

Fig. 14. D. wilsoni from Weddell seal. Transverse section of muscular 
system. 

Fig. 15. Glandicephalus antarcticus from Ross seal. Transverse section 
of muscular system, showing network formed by transverse and dorso- 
ventral musculature, enclosing in its ' meshes ' the longitudinal muscle- 
fibres. 

Fig. 16. Glandicephalus perfoliatus from Weddell seal. Transverse section 
of muscular system, showing similar structure as in G. antarcticus. Upper 
part of cortical parenchyma has not been depicted. 

Fig. 17. Baylisia baylisi from Crabeater seal. Transverse section of 
muscular system. 

Fig. 18. Baylisiella tecta from Elephant seal. Transverse section of 
muscular system, showing its peculiar structure and the powerfully 
developed cortical excretory system. Only a part of cortical paren- 
chyma and vitelline glands have been drawn. 






PLATE 11 



v A 



fliilii 








vq. 



T 

Lm. 
12 






-l.m. 



dvm 







. !•">■ vq. 
t.m. ^ 

II 







13 



MB* 



V-r^ 






M /*? ' 



( ■>/>* 07 



JVV< '/Y^ V"" 





S^Hhi 





^©® ©© rap 



dvm. 







1 /'••' * /' ; . ' 




17 









V 
& 

&W.1 



THE CESTODES OF SEALS FROM THE ANTARCTIC 133 

sections, shown in the Table No. 2, are more constant for a given species, although 
the numbers of the testes seen in the sagittal plane may differ in the same individual 
according to the shape of the segment. This was discovered in D. lasheyi, which 
possesses two types of gravid segments : squarish and elongate. In one of the squarish 
segments there are about 10, in the sagittal plane, and in the elongate segments 17 
testes. It seems that the size of the segment has no effect on the number of testes. 
In Diphyllobothrium scoticum, for example, where the variations in size of the segments 
are enormous, but the shape is the same, the number of testes in the sagittal plane of 
the small and the large segments is the same. 

The longitudinal muscular coat, examined in transverse section, and its relation to 
the transverse and dorso- ventral muscles also gives very clear specific differentiation. 
The thickness of the longitudinal muscular coat and the arrangements of the muscular 
bundles are characteristic for each species. The numerical differences in the thickness 
of the muscular coat have been given in Table No. 2. The morphological differences 
are shown in Plate 11, figs. 10-18. 

Examination of the musculature has been made in transverse sections of the hinder 
part of the segment, between the ovary and the first uterine coils. This part of the 
segment is not affected by such factors as the pressure of the uterus filled with eggs 
or histological changes of the ovary, which may interfere with or alter the position 
of the muscles. The relation of the transverse musculature to the longitudinal is 
also different in many cases (PL n, figs. 10-18). The transverse musculature in 
D. quadratum is so feebly developed that the separate muscle-fibres might be over- 
looked in the serial sections. A similar feeble development of the transverse muscula- 
ture occurs in D. mobile, D. wilsoni, and D. lasheyi, although in these the single 
muscle-fibres are much more distinct. In the rest of the species, namely, D. scoticum, 
Baylisia baylisi, and Baylisiella tecta, the transverse and longitudinal muscular coats 
are very powerfully developed. Moreover, the longitudinal muscles in B. tecta are 
very characteristic. 

In Glandicephalus antarcticus and Diphyllobothrium perfoliatum the transverse and 
the dorso-ventral muscular system forms a kind of network containing the longi- 
tudinal muscular fibres in its meshes. Because of the similarities in the musculature 
of these species, together with a few more points of resemblance in their anatomy, 
D. perfoliatum has here been transferred to the genus Glandicephalus Fuhrmann 
(1920). 

SYSTEMATIC NOTES 

i. DIPHYLLOBOTHRIUM Cobbold (1858) 
This genus is represented in the Antarctic seals by five species. 

Diphyllobothrium lashleyi (Leiper & Atkinson, 1914) 

[PL. 10, FIG. i; PL. II, FIG. 10; PL. 12, FIGS. 1 9-24] 

Dibothriocephalus lashleyi Leiper & Atkinson, 1914. 
Diphyllobothrium lashleyi Meggitt, 1924. 
Dibothriocephalus archer i Leiper & Atkinson, 19 14. 



i 34 THE CESTODES OF SEALS FROM THE ANTARCTIC 

Diphyllobothrium archeri Meggitt, 1924. 

Host : Weddell seal (Leptonychotes weddelli) . 

Locality : Debenham Islands ; Deception Island ; Melchior Archipelago. 

This tapeworm has been found in large numbers in the intestine of seven Weddell 
seals. Leiper & Atkinson (19 14) give its maximum length as 4 cm. and Johnston 
(1937) 22 mm. The variation in the length of the body as shown by the new material 
is much greater, from 4 cm. to 19-5 cm. The width is constant at about 3 mm. 

The segments are not overlapping and are variable in shape. Those in front are 
squarish with curved lateral edges ; those in the hinder part of the body become elon- 
gate. Both kinds of segment may be fully gravid. The size of the first squarish 
segment bearing eggs is, in mounted specimens, 1 mm. in length and 2 mm. in width. 
The elongate segments are about 7 mm. in length and 3 mm. in width. The terminal 
segment is usually slightly tapering and rounded at the end. 

The surface of the body in preserved specimens bears transverse furrows, caused 
probably by the fixative. 

The parenchyma is very loose and delicate, and mounted specimens are very trans- 
parent, showing the internal structure. 

The neck is well developed in fully extended specimens and marked off from the rest 
of the body. In specimens slightly contracted it is not distinguishable. As a con- 
sequence it varies from 450 /x to 2-5 mm. in length. 

The scolex, sharply marked off from the body, is also variable in shape; it is 
globular or oval and possesses internal grandular cells, though this was not previously 
known. The size of the scolex in mounted specimens is from 1 mm. to 1-5 mm. in 
length and 0-9 mm. to 1-3 mm. in width. 

The genital openings are median, except the uterine pores which alternate in an 
irregular manner. The genital atrium is surrounded with strongly developed papillae. 

In optical view in whole preparations the cirrus-sac appears to be spherical. 
In sagittal section its length is about 200 /x and the height (antero-posterior) about 
93 jit. The cirrus of the specimens examined was usually protruded. The cuticle of 
the area around the male genital pore is provided with radiating ridges. In the 
segments examined the vesicula seminalis is in a straight line with the cirrus-sac or 
inclined to it at a slight angle, and in sagittal section it is about 69 /x long and 55 fx 
wide, with walls about 17 /x thick. 

The testes are arranged in a single layer, about 60 on each side of the segment. 
They are about 53 /x to 93 /x in diameter. In the squarish segments there are, in 
transverse section, 2-6 testes each side, and, in sagittal section, 10 testes. In the 
elongate segments the number of testes in transverse section is 3 on each side and in 
sagittal, 15-17- 

The uterine opening is situated about 116 /x from the cirrus-sac. The uterus forms 
compact, though not very distinct, coils, and in the elongate segments is club-shaped, 
tapering posteriorly, and is irregular in outline. In the squarish proglottids the uterus 
is confluent with the cirrus-sac, reaching its level and alternating with it to its right 
or left side rather irregularly. 

The vagina and the male pore open side by side into the common genital atrium. 

The ovary has a compact or reticular structure. It differs with the shape of the 



PLATE 12, FIGS. 19-24 

(For list of abbreviations see Plate 10) 
Diphyllobothrium lashleyi from Weddell seal 

Fig. 19. Scolex. 

Fig. 20. Gravid, not fully shaped segment. 

Fig. 21. Gravid, fully developed elongate segment. 

Fig. 22. Transverse section of segment. 

Fig. 23, Sagittal section of segment. 

Fig. 24. Sagittal section of scolex, showing cephalic glands. 



PLATE 12 




THE CESTODES OF SEALS FROM THE ANTARCTIC 135 

segment. In the squarish segment it is more compactly built and oval in shape ; in the 
elongate segments its structure is reticular and more diffuse. 

The eggs are 53-56 /x x 40-44 p. 

The vitelline glands, composed of large cells, are irregular with rounded lobes. 
They are sometimes slightly confluent in the anterior parts of the segment and much 
more so in the posterior region. There is an area free from vitelline glands around 
the genital complex. The products of the glands are collected in special ducts running 
through the cortical parenchyma. It seems that there occur additional collectors, 
transferring the yolk to the main yolk reservoir which is situated in the central part 
of the segment below the uterus. The reservoir is filled with large yolk cells, olive- 
greenish in colour. The size of the vitelline glands is 33 fx to 50 jjl in diameter. 

The longitudinal muscles form a coat, 20 jjl thick in transverse section, composed of 
irregularly distributed bundles. The dorso-ventral musculature is feebly developed. 

The excretory system consists of two main stems in the medullary parenchyma and 
some 28 vessels occurring in the cortical part of the segment , as seen in transverse section . 

Leiper & Atkinson (1914) described D. archeri from the Weddell seal as a separate 
species. Comparison of the type specimens of D. archeri and D. lashleyi with the new 
material leads to the belief that D. archeri is a synonym of D. lashleyi. Both authors 
dealt with specimens not fully developed and having squarish segments, but the 
anatomical features of the two species, as seen in serial sections of the type specimens, 
are closely similar. The structure of the longitudinal muscles, cirrus-sac, and the 
number and arrangements of testes are identical. 

Diphyllobothrium mobile (Rennie & Reid, 1912) 

[PL. 10, FIG. 2; PL. II, FIG. Ii; PL. 13, FIGS. 25-31] 

Dibothriocephalus mobilis Rennie & Reid, 191 2. 

Diphyllobothrium mobile Meggitt, 1924. 

Diphyllobothrium wilsoni Railliet & Henry, 191 2. 

Dibothriocephalus coatsi Leiper & Atkinson, 1914 {nee Rennie & Reid, 1912). 

Host : Weddell seal (Leptonychotes weddelli) . 

Locality : Debenham Islands. 

This species is recorded from seven Weddell seals, twice in company with Glandi- 
cephalus perfoliatus. The infection was in most cases a mass infection. 

The length of the body of the specimens examined varied from 2-3 mm. to 14 mm., 
and the width from 345 /x to 510 /*. 

The strobila is composed of about 14 segments, which in the anterior part of the 
strobila are wider than they are long. The terminal segment is usually oval. 1 

The scolex is 675 /x-825 /x in length and 345 /u-510 /x in width. Longitudinal serial 
sections show that it contains glandular tissue. 

The neck is little more than an unsegmented part of the scolex, 300 t/, in length and 
495 /x in width. 

1 Beside the normally-developed segments, one abnormal strobila has been found in the writer's 
material. In it, the segments are split into two parts. The left-hand portion forms a kind of cul-de-sac 
and possesses testes and vitelline glands. The right part, which is a continuation of the strobila, contains 
eggs. 



136 THE CESTODES OF SEALS FROM THE ANTARCTIC 

The genital rudiments occur in the immediate vicinity of the scolex. The specimens 
of 2 *3 mm. in length have distinctly visible genital anlagen and well-separated testes. 

The length of the cirrus-sac, measured in sagittal section, is about 112 p and its 
height 66 jjl. Some very thin muscular fibres attach the proximal part of the cirrus-sac 
to the dorsal wall of the segment. 

The vesicula seminalis measures, in sagittal section, about 83 \l by 15 /x ; its walls 
are about 10 p thick and it is in a straight line with the cirrus-sac. 

The testes seem to occur only in the anterior part of the segment and are from 
22 to 44 in number. There are 5 testes on each side in the transverse, and 6 to 7 in 
the sagittal, sections of the segment. They are 66 [i to 99 /x in diameter. 

The uterine openings alternate irregularly. The uterus forms a compact mass of 
coils. In the anterior part of the body it is situated below the cirrus-sac, and in 
segments in the hinder part of the strobila its coils surround the male copulatory organ. 

The vagina opens into the common genital opening in the vicinity of the male 
opening. 

The ovary forms two more or less elongate-oval wings. 

The eggs measure 56-60 fi by 40-43 /x. 

The vitelline glands, about 33 \l in diameter, are arranged in two separate lateral fields 
in each segment. A narrow transverse space free of vitelline glands is distinctly visible 
in the anterior part of the segment in whole preparations as well as in serial sections. 

The longitudinal musculature is very feebly developed. It forms a coat about 4 \i 
thick, composed of single, barely visible, fibres. 

The excretory system has not been detected, probably owing to contraction due to 
the fixative. 

Diphyllobothrium quadratum (Linstow, 1892) 

[PL. 10, FIG. 3; PL. II, FIG. 12; PL. 14, FIGS. 32-36] 

Bothriocephalus quadratus Linstow, 1892. 
Dibothriocephalus quadratus Zschokke, 1903. 
Diphyllobothrium quadratum Railliet & Henry, 191 2. 
Cordicephalus quadratus Ward, McLeod & Stewart, 1947. 
Dibothriocephalus coatsi Rennie & Reid, 191 2. 
Bothriocephalus coatsi Fuhrmann, 1920. 
Dibothriocephalus resimum Railliet & Henry, 191 2. 

Host : Leopard seal (Hydrurga leptonyx) . 

Locality: Galindez Island, Argentine Islands; Debenham Islands; Horseshoe Island and 
Sandefjord Harbour, Coronation Island. 

This species is recorded from five Leopard seals, some of which were very heavily 
infested. 

The length of the body is from 4 to 12 cm. and the width about 4-5 mm. The 
specimens obtained from the mass infested hosts averaged about 4 cm. in length. In 
horizontal serial sections the lateral margins of the body seem to have a villous 
character. 

The segments are square, about 1-5 mm. long and 1 to 4-5 mm. in width. Their 
lateral edges in the hinder part of the strobila are slightly convex. The terminal 
segment is oval. 



PLATE 13, FIGS. 25-31 

(For list of abbreviations see Plate 10) 
Diphyllobothrium mobile from Weddell seal 

Fig. 25. Malformation of strobila. 

Fig. 26. Young specimen with genital anlagen. 

Fig. 27 (a). Sagittal section of scolex, showing glandular tissue, (b) 
Glandular cells enlarged. 

Fig. 28. Terminal gravid segment. 

Fig. 29. Transverse section of segment. 

Fig. 30. Sagittal section of segment, showing terminal part of uterus. 

Fig. 31. Sagittal section of male and female openings. 



PLATE 13 



\l ''b' 6 27 




Figs. 25-31 



S&Ti7 t 






PLATE 14, FIGS. 32-36 

(For list of abbreviations see Plate 10) 
Diphyllobothrium quadratum from Leopard seal 

Fig. 32. Scolex. 

Fig. 33. Gravid segment. 

Fig. 34. Horizontal section of segment, showing structure of ovary 

Fig. 35. Transverse section of segment. 

Fig. 36. Sagittal section of segment. 



PLATE 14 




ma m -J 1 t 







05 mm. 






\MU^\Mi^^^ 








. ^-<^. ^W - 



35 








:«ftgi 



•-< « 



St. O o - T J J - 

; t ', ;>0 3 50 . 









i 



3 a r © ooo ©„;; o«i 



!.© &V«°« '-^o , o^r?©? 



0-5 mm. 



-v.q 



33 



THE CESTODES OF SEALS FROM THE ANTARCTIC 137 

The scolex of the mounted specimens is 1 to i-8 mm. in length and 960 fi to 1-4 mm. 
in width, more or less ovoid in shape and possesses internal glandular tissue. 

The neck although short is recognizable and measures from about 450 /x to 1-5 mm. 
in length and from 587 to 870 1* in width. 

The genital rudiments occur in the segment next behind the neck. In whole 
preparations the cirrus-sac appears to be spherical ; in sagittal section its length is 
about 300 [A and height 60 ju,. 

The vesicula seminalis is connected with the cirrus-sac almost along the same 
axis and is 120 ju, in length and 105 /x in width. The walls are about 17 (jl thick. The 
almost spherical shape of the organ is sometimes rendered more or less irregular, 
probably by the fixative used. 

There are about 340 testes in each segment, 170 on each side and confluent in its 
anterior part. They are arranged in a single layer, though this is not very regular 
in some cases, probably owing to contraction. This was observed in the small 4 cm. 
long specimens, where some of the testes were arranged in two planes. There are 12 
to 16 testes on each side in the transverse, and 12 to 16 in the sagittal, section. They 
are not very regular in shape and measure about 65 jjl by 78 /x. 

The vas deferens is situated dorsally in the segment and forms numerous coils. 

The uterine openings alternate irregularly. In the hinder region of the strobila the 
uterus is converted into an irregular sac filled with eggs. The anterior part of the 
uterus surrounds the cirrus-sac which, in the gravid posterior segments, is hardly visible . 

The ovary forms two wings, which surround the lower coils of the uterus. The 
thin-shelled and operculate eggs are 56 /x by 43 \i. 

The vitelline glands are very numerous, thickly arranged, irregularly lobed, and 
measure 50 \l by 33 ^. They obscure all other organs of the genital complex except 
the uterus and the cirrus-sac. 

The longitudinal muscles, not very well developed, form a coat about 33 tt thick. 
They are composed of single bundles of fibres. 

The transverse and dorso-ventral musculature is composed of single fibres running 
in these two planes. 

The excretory system comprises two main vessels in the medullary parenchyma 
and about 34 trunks in the cortical part of the segment in transverse section. 

Diphyllobothrium scoticum (Rennie & Reid, 1912) 
[Pl. 10, fig. 4; Pl. 11, fig. 13; Pl. 15, figs. 37-43] 

Dibothriocephalns scoticus Rennie & Reid, 191 2. 
Diphyllobothrium scoticum Meggitt, 1924. 
Dibothriocephalus pygoscelis Rennie & Reid, 19 12. 

Host: Leopard seal (Hydrurga leptonyx). 1 

Locality: Debenham Islands. 

This species has been found in four Leopard seals. The number of worms per host 
varied from 2 to 14. 

1 Baylis (in Hamilton, 1934) assigned to Diphyllobothrium scoticum some Cestodes from the intestine of 
Otaria byronia, but the identification was only provisional and has not yet been confirmed. 
zoo. 1. 7. s 



138 THE CESTODES OF SEALS FROM THE ANTARCTIC 

The body is much longer than recorded by Fuhrmann (1920) and Johnston (1937) 
and ranges in the specimens examined from 52 cm. to 130 cm., with a corresponding 
width of 0-5 cm. to i-8 cm. The elliptical scolex is about 3-5 mm. in length and 2 mm. 
in width. No glandular tissue has been found in this organ. 

The neck is about 495 fx in length and 825 /x in width. The figures for scolex and 
neck have been taken from a mounted specimen 52 cm. long. 

The segments are shorter than wide and have convex lateral edges. They are 
tapering in the posterior part of the body, and the terminal segment in small speci- 
mens is ovoid. The posterior lateral edge of the segment seems to have a semicircular 
thickening (PL 15, fig. 43). The gravid segments are 5 to 8 mm. in length and 1-5 cm. 
to i-8 cm. in width. 

The genital rudiments occur in the first segment behind the neck. The male 
genital openings are surrounded by numerous papillae, radially arranged, and are 
sometimes bordered with a semi-lunar furrow. 

In sagittal section the cirrus-sac measures about 231 fi in length and 142 /z in height. 
In the vicinity of the cirrus-sac, plainly visible in the sagittal section, occur spherical 
cells, probably of a glandular character. 

The vesicula seminalis is situated in the same main axis, as a continuation of the 
cirrus-sac. It is about 285 ^ in length and 180 /a in width, with walls about 17 fi thick. 

The vas deferens runs dor sally in numerous coils. 

There are about 600 testes, disposed in two separate fields about 300 on each side, 
arranged in a single layer and measure about 150-210 \l by 150 [i ; they are not con- 
fluent in the anterior part of the segment. The number of testes in sagittal section 
amounts to 25. It seems that there is no difference in the number of testes in sagittal 
section as between the large and the small gravid specimens. The number of testes 
counted in transverse section amounts to 14-15 on each side. 

The uterine openings alternate irregularly, and are situated in a transverse groove. 
The terminal part of the uterus is modified to form a thick- walled pocket, lined with 
a villous tissue. This modification of the terminal uterine duct is typical for the 
species. The uterus is not of a 'rosette' type but forms spiral coils, more or less 
distinct, 5-12 in number on each side. 

The ovary is reticular and irregularly palm-shaped. The eggs, some provided with 
a boss, are 76-79 //, by 56 \x. 

The vitelline glands, composed of small cells, are fairly large, about 60-105 ju, in 
diameter. They are confluent in the anterior part of the segment, leaving a free area 
around the genital opening. 

The longitudinal, transverse, and dorso-ventral musculature is very well developed. 
The longitudinal muscles form a coat 150 /x thick measured in transverse section, 
composed of numerous fibres, collected in not very distinct bundles. They are thickly 
arranged in the upper parts of the muscular coat, gradually becoming less dense 
towards the middle of the segment. 

The excretory system consists of 2, not always distinctly visible, trunks in the 
medullary parenchyma and about 20 in the cortical parenchyma. It is, however, not 
always possible to be sure of the number of cortical excretory vessels as the vessels 
may contract as a result of fixation and may not be distinct in serial sections. 



PLATE 15, FIGS. 37-43 

(For list of abbreviations see Plate 10) 
Diphyllobothrium scoticum from Leopard seal 

Fig. 37. Gravid segment of small specimen. 

Fig. 38. Modification of terminal part of uterus, seen in transverse 
section. 

Fig. 39. Scolex. 

Fig. 40. Gravid segment of large specimen. Part of cortical parenchyma 
has been removed, showing distribution of testes and terminal modifica- 
tion of uterus. 

Fig. 41. Transverse section of segment. 

Fig. 42. Sagittal section of segment with male and female openings. 

Fig. 43. Outline of posterior edges of segment, in sagittal section. 



W>— I 




37 





goo o o8o° 'ogoo 

OoO°o |ooOogoo 

&°o°o0 o 0000 
%%o §o o 0o0 o 

OcPonO QOO°oOO 
°nOoo°OOo < 



Voo° o% o ■ 



j O ^' ^ 

o oc I o c 








39 




40 




N^-1- 









v!d. 




42 



Figs. 37-43 



THE CESTODES OF SEALS FROM THE ANTARCTIC 139 

Diphyllobothrium wilsoni (Shipley, 1907) 

[PL. 10, FIG. 5; PL. II, FIG. 14; PL. l6, FIGS. 44-50] 

Dibothriocephahis wilsoni Shipley, 1907. 

Dibothriocephahis scotti Shipley, 1907. 

Diphyllobothrium scotti Meggitt, 1924. 

Dibothriocephahis mobilis Leiper & Atkinson, 1915 (nee Rennie & Reid, 1912). 

Host : Weddell seal (Leptonychotes weddelli) and Leopard seal (Hydrurga leptonyx) . 

Locality: Deception Island; Beascochea Bay; Debenham Islands; Melchior Archipelago; 
Argentine Is. (Galindez I.) and South Sandwich. 

The species was found in eight Weddell seals and six Leopard seals. 

The length of the body varies from about 1 cm. to 5 cm. with a maximum width 
of 3 mm. 

The scolex is variable in size and shape, probably the result of contraction due to 
fixation; its length is about 825 /x and the width from 450 /x to 1 mm. The presence 
of glandular tissue in the scolex has been confirmed, using Ehrlich's haematoxylin 
and erythrosin. 

The neck is very short and often impossible to distinguish, 375 ju, in length and 
about 240 jjl in width. 

The first segments are shorter than wide, becoming gradually squarish, and then 
longer than wide. The terminal segment is usually oval or elongate-oval. 

The genital rudiments occur in the first segment behind the scolex. 

The cirrus-sac is 106-165 /"- l° n g an d 83-99 P high m sagittal section. It is attached 
in its hinder part, with numerous well-developed muscular fibres, to the dorsal wall 
of the segment. These structures are plainly visible in a sagittal section 10 /x thick. 
The contracted cirrus seems to be coiled spirally. When everted the cirrus is about 
44 ijl to 59 fx in length. 

The vesicula seminalis is about 92 /x in length and 56 /x in width, with the walls 
about 20 fi thick. 

The vas deferens, filled with sperm, runs dorsally and is irregularly coiled. 

There are about 150 testes, 75 on each side of the segment, arranged in a single 
layer and very closely distributed. They are confluent in the anterior part of the 
segment, irregular in shape, with slightly lobed outlines. There are 8 testes on each 
side in the transverse and 6 to 8 in the sagittal section ; their dimensions are about 
132 [jl by 99 jjl. Histologically, as compared with other species, the testes seem to be 
more compact and stain more intensively with Ehrlich's haematoxylin. This may, 
however, be due to the number of spermatozoa in the testicular tissue absorbing more 
of the stain. 

The uterine openings are irregularly alternating. The uterus forms an irregular 
spherical sac filled with eggs and surrounds the cirrus-sac on both sides. 

The vagina opens close to the male genital opening in the common genital atrium. 

The eggs are 50 /x by 40 /x. 

The ovary is irregular in outline or kidney-shaped with more or less lobated edges. 
It is composed of large egg cells. 

The vitelline glands, measuring 66 /x by 50 /x, are irregularly spherical and somewhat 



140 



THE CESTODES OF SEALS FROM THE ANTARCTIC 



amoeboid, very numerous, and strongly developed. They form thick uninterrupted 
layers from segment to segment, covering all the internal organs, except the uterus. 

The longitudinal musculature is not very strongly developed. It forms a coat 
composed of minute bundles about 17 /x thick in transverse section. 

The central, medullary excretory system consists of two main stems not very 
easily distinguishable. The cortical excretory system possesses approximately 14 
stems, fairly large in diameter, running among the vitelline glands. 

It seems from a comparison of the small specimens of D. wilsoni and the type 
specimens of D. scotti, described by Shipley (1907), with the newly-collected material 
from the Leopard seal, that D. scotti is a synonym of D. wilsoni. The comparison in 
Table 3 of Fuhrmann's data (1920) with the figures recorded from the present 

Table No. 3 
Comparison of D. wilsoni and D. scotti 





Fuhrmann 


{1920) 


Writer's material 




D. wilsoni 


D. scotti 


D. wilsoni 


Body: length ... 


10 mm. 


9 cm. 


5 cm. 


width 






1-7 mm. 


2 mm. 


1-5-3 mm. 


Scolex: length 






85O /Lt 


500-900 fi 


825 fi-i mm. 


width 






45o^ 


700 fX 


450 )L4-i mm. 


Neck: length 






? 


short 


375/* 


width 






? 


? 


240 /X 


Cirrus-sac: length 






140^ 


150^ 


106-165 (i 


width 






? 


? 


83-99 p 


Vesicula seminalis, diameter . 


80 (X 


80 n 


92X56 /Lt 


No. of testes: 








transverse section .... 


6-9 


6 


8 


sagittal section ..... 


6 


6-10 


6-8 


Eggs, diameter ..... 


60 X 36 fi 


64 X 4O fX 


50X40/U 


No. of cortical excretory vessels 


14 


12 


H 


Thickness of longitudinal muscle-layer 


12 /Li 


14-18 fi 


17 n 



material shows that there is practically no difference between the two so-called species, 
except in the size of the body, and it is this which seems to have misled previous 
authors. The small, mature specimen of D. wilsoni might be considered a dwarf form, 
caused by mass infection and consequent unfavourable living conditions. In the 
Leopard seal, where the infection is not so heavy, D. wilsoni reaches a relatively large 
size. 

2. GLANDICEPHALUS Fuhrmann 1920 

Glandicephalus antarcticus (Baird, 1853) 

[Pl. 10, fig. 6; Pl. 11, fig. 15; Pl. 17, figs. 51-53] 

Bothriocephalus antarcticus Baird, 1853. 
Dibothrium antarcticum Diesing, 1863. 
Diplogonoporus antarcticus Liihe, 1899. 
Dibothriocephalus antarcticus Shipley, 1907. 
Diphyllobothrium antarcticum Railliet & Henry, 191 2. 
Glandicephalus antarcticus Fuhrmann, 1920. 
Host: Ross seal (Ommatophoca rossi). 



PLATE 16, FIGS. 44-50 

(For list of abbreviations see Plate 1 0) 
Diphyllobothrium wilsoni from Weddell seal 

Figs. 44-46. Scolex in different stages of contraction. 

Fig. 47. Gravid segment. 

Fig. 48. Terminal segment with double set of gravid genital organs. 

Fig. 49. Transverse section of segment. 

Fig. 50. Sagittal section of segment. 



PLATE 16 





Q-5 mm. , 



Figs. 44-50 



PLATE 17, FIGS. 51-53 

(For list of abbreviations see Plate 10) 
Glandicephalus antarcticus from Ross seal 

Fig. 51. Scolex. 

Fig. 52. Sagittal section of several segments. 

Fig. 53. Transverse section ol segment. 



PLATE 17 




I mm. 




53 



).i.7. 



THE CESTODES OF SEALS FROM THE ANTARCTIC 141 

This species, described by Baird (1853) and re-described by Fuhrmann (1920), 
was collected by the Ross's Antarctic Expedition. To give a complete picture of the 
Pseudophyllidean Cestodes occurring in Antarctic seals, however, the type-specimens 
have been re-described to show their generic relationship with Glandicephalus 
perfoliatus. 

The specimens re-examined were about 10 cm. in length and 7 mm. in width. The 
strobila is markedly imbricate. 

The scolex, 3 mm. in length and 2 mm. in width, is provided with protuberances 
as depicted by Baird, though these are not distinct in all specimens. No glandular 
structure was discovered in sagittal serial sections stained with Ehrlich's haema- 
toxylin. 

The neck is about 2-35 mm. long and 750 /x in width. 

The cirrus-sac, in sagittal section, measured about 462 /x in length and 231 fx in height. 

The vesicula seminalis is about 132 fx by 155 jx, with walls about 10 [x thick. 

The testes are scattered in the medullary parenchyma and among the longitudinal 
musculature. They are arranged in irregular layers. 

The eggs are 43-50 \x by 33 \x and the vitelline glands are about 40 \x by 50 ll. 

The muscular system, as described and depicted by Fuhrmann (1920), is almost 
identical with that of G. perfoliatus. The longitudinal muscular coat is about 
450 [x thick. 

There are about 30 excretory vessels, counted in the transverse section of the 
cortical parenchyma. 

Glandicephalus perfoliatus (Railliet & Henry, 1912) n. comb. 

[PL. 10, FIG. 7; PL. II, FIG. l6; PL. l8, FIGS. 54-59] 

Diphyllobothrium perfoliatum Railliet & Henry, 191 2. 
Dibothriocephalus perfoliatus Fuhrmann, 1920. 
Diphyllobothrium clavatum Railliet & Henry, 191 2. 
Diphyllobothrium rufum Leiper & Atkinson, 19 14. 
Host : Weddell seal (Leptonychotes weddelli) . 

Locality: Debenham Islands; Stella Creek, Deception Island; Argentine Is. (Galindez I. and 
Winter I.) ; Beascochea Bay, Graham Land (Hut Cove, Hope Bay) ; and Palmer Archipelago 
(Port Lockroy, Wiencke Island) . 

This species was collected from thirteen Weddell seals. It occurs usually as a mass 
infection, mainly in the bile-duct, overhanging into the gut. The specimens examined 
were at different stages of maturity from 4 mm. to 20 cm. in length. The average 
length of the body ranges from 12 to 14 cm., and the minimum width is 7 mm. 

The strobila is differentiated into two distinct parts: the anterior, ivory white, 
amounting to about one-third of the total length, and the posterior, yellowish, 
increasing in width and tapering slightly at a small distance from the posterior end of 
the body. This differentiation of the strobila has not been observed in specimens of 
4 cm. or 5 cm. in length, in which the outline of the body is oval with a well-defined 
scolex. The surface of the strobila bears, a small distance behind the neck, charac- 
teristic strongly developed imbrications formed by the excessive development of the 
cortical part of the segment. 



i 4 2 THE CESTODES OF SEALS FROM THE ANTARCTIC 

The scolex is about 3-5 mm. in length, measured in formalin specimens, and 2 mm. 
in width. No glandular structure has been discovered in longitudinal sections of the 
organ. 

The neck is distinct and ranges from 2-5 mm. to 3 mm. in length and 1 mm. in 
width. 

The segments are very short. The length of a fully gravid proglottid, measured 
in a sagittal serial section, is about 400 jx. The terminal segment is small and bell- 
shaped, with a cone at the terminal part of its longitudinal axis. It contains normal 
genital organs and produces eggs. This peculiar shape of the segment may be the 
result of fixation. 

The genital openings are irregularly alternating and situated on the anterior 
surface of the imbrications. 

The length of the cirrus-sac, measured in a sagittal section, is about 264 it, and its 
width 188 ft. 

The pear-shaped vesicula seminalis is 148 /x long and 132 /x wide in sagittal 
section ; its walls are about 20 /x thick. 

There are about 100 testes in the fully gravid proglottid. They are more or less 
distributed in a single layer, which is more readily distinguishable in the lateral part 
of the segment. Near the centre, close to the uterine coils, the testes are arranged in 
irregular clumps. In transverse section there are about 14 testes on each side, and in 
the sagittal plane from 3 to 6 ; they measure 1 16-172 fi by 73-93 /x. 

The vagina opens immediately behind the male pore, slightly obliquely. 

The uterine openings are irregularly alternate on the left or right side of cirrus-sac, 
and the uterus forms an irregular sac with indistinct coils, filled with eggs. These, 
some of them provided with a boss, are about 60-66 /x by 50 /x. 

The ovary comprises two small wings tapering towards the lateral edge of the 
segment. 

The vitelline glands, distributed mainly in the anterior region of the imbrications 
of the segment, are very irregular in shape and size and measure about 70 /x by 40 /x. 

The longitudinal musculature of this species is very unusual and almost identical 
in structure with that of G. antarcticus (Baird). Together with the transverse and 
dorso-ventral muscular system, it is distributed throughout almost the whole medul- 
lary part of the segment. The single bundles of the longitudinal muscular system 
are separated by the fibres of the dorso-ventral and transverse musculature. 
Examined in transverse section the two lateral systems form a kind of square which 
encloses, at its centre, the fibres of the longitudinal muscles. The boundary between 
the cortical and medullary parenchyma, so characteristic of, and fairly easily dis- 
tinguishable in, most other species of Pseudophyllidean Cestodes, is not very distinct 
in G. perfoliatus, because of the network formed by the transverse and dorso-ventral 
musculature, through the meshes of which run the fibres of the longitudinal muscles. 
In the near vicinity of the testes, or of the excretory vessels, the muscular system is 
more diffuse. 

The central excretory system in the medullary parenchyma is composed of 2 vessels, 
which run an undulating course through the length of the body and are about 17 /x 
in diameter. The cortical excretory system seems to be composed of 16 vessels. 



PLATE 18, FIGS. 54-59 

{For list of abbreviations see Plate 10) 

Glandicephalus perfoliatus from Weddell seal 

Fig. 54. Young immature specimen. 

Fig. 55. Young specimen showing differentiation of strobila. 

Fig. 56. Portion of strobila showing internal structure, after removal of 
part of cortical parenchyma. 

Fig. 57. Sagittal section of segments with male and female openings. 

Fig. 58. Sagittal section of two posterior segments. 

Fig. 59. Transverse section of segment, with vitellaria in portion of 
imbrication. 



PLATE 18 





58 




at'U'UII/0,,^ 








V^ f 



56 





O-S mm. 



57 




wmimm 







59 
Figs. 54-59 



THE CESTODES OF SEALS FROM THE ANTARCTIC 143 

The two main nerves, 66 jjl by 44 ju, in transverse section, are situated outside the 
medullary excretory system. 

Comparison of this material with the type specimens of Glandicephalus antarcticus 
(Baird) reveals that, although there is no doubt that there are two distinct species, 
these have features in common that appear to warrant their being grouped together 
in a single genus distinct from Diphyllobothrium ; the name Glandicephalus Fuhrmann 
(1920) is available. It is true that this name was coined by Fuhrmann in consequence 
of his discovery of glandular tissue in the scolex of antarcticus and that no such 
tissue has been found in the scolex of perfoliatus ; but this character is not at all 
diagnostic of Glandicephalus and occurs in several species of Diphyllobothrium, e.g. 
D. lashleyi, D. mobile, and D. quadratum. Features common to both antarcticus and 
perfoliatus, but not found in any true Diphyllobothrium, are as follows : imbrication of 
the segments (less well developed in the first species) ; presence of a well-separated 
neck, and the arrangement of the musculature. In both species the transverse and 
dorso-ventral muscles form a kind of network spread in the medullary parenchyma, 
enclosing in the 'meshes' the fibres of the longitudinal muscles. Moreover, the 
vitelline glands are situated in the anterior region of the imbrications of both species. 
Finally, they seem to show a host specificity: G. antarcticus for Ommatophoca and 
G. perfoliatus for Leptonychotes weddelli. 

Leiper & Atkinson (1914) describe another species from the Weddell seal, namely, 
Diphyllobothrium rufum. Johnston (1937), who examined immature specimens which 
he thought to be D. rufum, was apparently inclined to believe that the species was 
probably a 'precocious form* of G. perfoliatus. Comparison of the type-specimens of 
D. rufum with G. perfoliatus reveals that the only differences are in the shorter neck 
and the presence of a 'notch' in the posterior margin of some of the segments. The 
neck of D. rufum, measured in sagittal section, is over 1 mm. in length. To judge from 
the published descriptions there would seem to be a difference in egg size also, for 
Leiper & Atkinson report the eggs of D. rufum to be 25 \i in diameter. This, however, 
appears to be a slip ; in the type specimens their dimensions are 59-66^ by 43-46 jit, 
which is identical with the size of the eggs in G. perfoliatus. Further, there are no 
differences in the structure of the genital apparatus or of the musculature. Diphyl- 
lobothrium rufum seems, therefore, to be a synonym of Glandicephalus perfoliatus. 

3. BAYLISIA gen. nov. 

Diagnosis: Large Cestodes in which the anterior part of the body is cylindrically 
modified. Scolex with cup-shaped bothria. Normal segmentation not distinct ; the 
body bears pseudosegmentation not corresponding to the individual sets of genital 
organs. 

Genital organs and their openings situated ventrally on both sides of the segment 
in double sets, regularly alternate in relation to the main axis of the body. 

Testes arranged in a single layer. Ovary ramified. Longitudinal muscles forming 
a thick coat. Excretory system situated in the cortical parenchyma. 

Type species : Baylisia baylisi sp. nov. 

Type host: Crabeater seal {Lobodon carcinophagus) . 



i 4 4 THE CESTODES OF SEALS FROM THE ANTARCTIC 

Baylisia baylisi sp. nov. 

[Pl. io, fig. 8; Pl. ii, fig. 17; Pl. 19, figs. 60-68] 

Host : Crabeater seal (Lobodon carcinophagus) . 
Locality : Deception Island ; Debenham Islands. 

This parasite has been found in two Crabeater seals. There were two complete 
worms, one 35 cm. in length and 1 cm. in width and a second 126 cm. by about 
8 mm., together with some fragments of strobila, varying from 15 cm. to 63 cm. in 
length and from 11 mm. to 15 mm. in width. 

The colour in formalin is ivory-white in the anterior part of the body, becoming 
brownish-grey in the posterior segments. The ivory-white part of the strobila, about 
2 cm. in length, is more or less cylindrical. 

There are two double furrows running laterally along the body. The central part 
of the strobila is convex along the main axis, marking the position of the uterus. 

The scolex has two cup-shaped bothria, and measures, in the specimen mounted in 
Canada, balsam about 900 /x long and 1-3 mm. broad. 

A neck seems not to be developed since segmentation starts immediately behind 
the scolex. 

The segmentation is very distinct, but apparently does not divide the body into 
single genital complexes, as happens in other tapeworms. The 'segments' are 2 cm. 
long and about 1-5 cm. in width, the terminal one being oval. 

The genital openings are situated ventrally on a segment in the longitudinal 
furrows, alternately left and right. There are about 30-40 double genital sets in one 
'segment', which alternate in relation to the main axis of the body, being arranged 
in 'zigzag', in contrast to Diplogonoporus, where they occur as two genital sets in the 
same transverse plane. In Baylisia baylisi one set of the genital organs only may be 
seen in transverse section. 

The cirrus-sac measures 750 /x long and 180 /x high in sagittal section. An irregularly 
coiled ductus ejaculatorius is situated inside the cirrus-sac. A cirrus has not been 
observed. 

The vesicula seminalis is situated slightly laterally but internally to the cirrus-sac, 
and is 198 lx long and 99 /u. wide in sagittal section ; its walls are about 33 ^ to 50 /x 
thick. The internal surface of the walls seems to have a villous structure. 

Beside the normally developed cirrus-sac mentioned above, one abnormality has 
been noted. Two cirrus-sacs were joined together, with a common opening, but each 
with a separate vesicula seminalis (PL 20, fig. 65). 

In each segment there are about 36 testes arranged in a single layer and flattened 
antero-posteriorly. The number of testes in transverse section amounts to about 
18 on each side, and there is 1 testis in sagittal section. The testes measure 165 /x 
in diameter in transverse section and 40 /x in the sagittal plane. They are elongate- 
oval, 116 [jl by 40 fju in horizontal section, with the longer axis transverse to the main 
line of the segment. 

The vagina opens into the genital atrium with the male opening and runs ventrally, 
but is transverse to the terminal part of the uterus. 



PLATE 19, FIGS. 60-68 

(For list of abbreviations see Plate 10) 
Baylisia baylisi from Crabeater seal 

Fig. 6o. Scolex. 

Fig. 6i. Sagittal section of fused cirrus-sac, with common genital duct 
and opening. 

Fig. 62. Transverse section of part of male and female genital apparatus. 

Fig. 63. Transverse section of segment, showing structure of ovary and 
genital ducts. 

Fig. 64. Sagittal section of part of segment, showing arrangement of 
ovaries and uterine coils. 

Fig. 65. Transverse section of male and female ducts (enlarged). 

Fig. 66. Transverse section of segment. 

Fig. 67. Sagittal section of segment, showing male copulatory organs. 

Fig. 68. Horizontal section of part of strobila, showing arrangement of 
genital organs. 



PLATE 19 



0> +r. % 
















6 4 



^•^ * 62 











o cb. 



63 



66 




65 




"ow 



60 





W(/^I 






I .1 



/ ■ 

1 



WfflWvi^MM 



68 




).i. 7. 



Figs. 60-68 
u 



THE CESTODES OF SEALS FROM THE ANTARCTIC 145 

The uterine openings on both sides are situated nearer the median line than the 
genital openings and lie posteriorly and obliquely to the cirrus-sac. The uterus itself 
comprises a few horizontal coils in the central part of the segment. 

The ovary is very characteristic and is composed of one solid compact central part, 
giving off a system of branches and ramifications among the uterine coils; in 
sagittal section it is V-shaped. 

The eggs are 66 /x by 46 xx. 

The musculature is very well developed. The longitudinal muscles form, in trans- 
verse section, a continuous coat about 450 /x thick, composed of numerous bundles. 
The transverse muscles are also well developed but the dorso-ventral musculature is 
much weaker. 

The excretory system seems to occur only in the cortical parenchyma, and is 
composed of about 76 vessels, visible in transverse section. Their transverse diameters 
differ considerably. This cortical excretory system is situated near the surface of the 
segment and runs among the vitelline glands. 

The vitelline glands, 43 /x by 20 xx, are composed of tiny cells and form a continuous 
layer, interrupted by the excretory system. They are more or less irregularly elongate 
in sagittal section, and elongate-oval in the horizontal plane. 

'Calcareous' bodies are very numerous, 17 /x by 26 /x in diameter, and occur in the 
cortical and medullary parenchyma. They have also been noticed among the 
longitudinal muscles. 

4. BAYLISIELLA gen. nov. 

Diagnosis: Pseudophyllidean Cestodes with the scolex bearing two strongly de- 
veloped bothria, modified in the anterior part in a f oliaceous lamella. The thick strobila, 
composed of very short and broad segments, is tapering posteriorly. The testes, 
arranged in two or three layers in the medullary parenchyma, have a tendency to 
ascend dorsally in relation to the uterus. The longitudinal muscles form, in transverse 
section, club-shaped or elongately oval bundles, some of them collected in pyramids, 
separated by the excretory vessels. Excretory system occurs in the cortical 
parenchyma. 

Type species : Baylisiella tecta (Linstow) . 

Type host: Elephant seal (Macrorhinus leoninus). 

Baylisiella tecta (Linstow, 1892) 

[PL. 10, FIG. 9; PL. II, FIG. l8; PL. 20, FIGS. 69-72] 

Bothriocephalus tectus Linstow, 1892. 
Dibothriocephalus tectus Zschokke, 1903. 
Diphyllobothrium tectum Meggitt, 1924. 
Cordicephalus tectus Wardle, McLeod, & Stewart, 1947. 

Host: Elephant seal (Macrorhinus leoninus). 

Locality: Bay of Isles, South Georgia. 

This species was found in two Elephant seals ; there were three entire worms in one 
host and two in the other, one specimen being headless. 

ZOO. I. 7. U2 



146 THE CESTODES OF SEALS FROM THE ANTARCTIC 

The thick belt-shaped strobila, composed of very short indistinct segments, is 32 
cm. in length and 2 cm. in width. 

The previous descriptions of Linstow (1892) and Fuhrmann (1920) were based on 
headless specimens. 

The scolex, not previously described, is deeply embedded in the intestinal tissue 
and very characteristic in shape. It possesses two powerful bothria and a complicated 
lamellar structure of their upper part, which recalls the scolex of Pyramicocephalus 
Monticelli 1890. The ' cauliflower ' lamellar structure seems to give rise to two lateral 
lamellar flaps and there is a similar differentiation on the top of the scolex. The 
length of the scolex is 8 mm. and the width 5 mm., and the transverse diameter across 
the 'flap' is 12 mm. 

The neck seems to be very short or not-existent. 

The segments are very short, about 165 /x-200 /x in length. 

The genital pores are situated in a common recess, provided with numerous papillae. 

The cirrus-sac, measured in sagittal section, is about 450 [x in length and 150 /x in 
height. 

The vesicula seminalis, situated in the same axis as the cirrus-sac, is 195 /x in 
length and 180 fx in width ; its walls are about 30 /x thick. 

The vas deferens runs dorsally in numerous coils. 

The testes are distributed in 2 or 3 layers, some of them ascending almost dorsally 
to the uterus and close to it. There are about 45 testes on each side in transverse, 
and 2 or 3 in sagittal, sections. They measure about 136 /x by 86 fx. 

The uterine openings are situated a little below the cirrus-sac on the right side. 
The uterus comprises a few irregular transverse coils. 

The vagina opens in the vicinity of the cirrus-sac on its right side. 

The ovary is bilobed and elongate. 

The eggs with 3 [x thick shells, measure 59-66 /x by 46 /x and are thickened opposite 
to the operculum. 

The vitelline glands, 66 fx by 26 /x, and arranged in a very thick layer, are very 
numerous, spherical or oval in transverse section. They form a compact mass of 
glands in the cortical parenchyma. 

The excretory system seems to occur in the cortical part of the segment. It is very 
strongly developed and runs through the longitudinal muscular system with numerous 
transverse anastomoses. In transverse section about 108 excretory canals have been 
counted. 

The musculature is exceedingly well developed. The longitudinal muscles are 
collected in large irregular bundles, which, in transverse section, appear club-shaped 
or elongate-oval. The bundles are often separated into distinct groups between 
which run excretory canals and dorso- ventral muscle-fibres. The thickness of the 
longitudinal muscular coat is about 555 /x. There is also a layer of longitudinal 
muscles situated in the subcuticular region of the segment, externally to the vitelline 
glands. This additional layer is composed of scattered, rather thick, individual fibres 
or very small bundles of fibres. The transverse muscles are also very strongly 
developed. 

This species represents a new genus, Baylisiella, distinguished from Bothriocephalus 



PLATE 20, FIGS. 69-72 

{For list of abbreviations see Plate 10) 
Baylisiella tecta from Elephant seal 

Fig. 69. Transverse section of segment. 
Fig. 70. Transverse section of male apparatus. 
Fig. 71. Sagittal section of segment. 
Fig. 72. Scolex. 



PLATE 20 










smez? 






^ 






69 




Figs. 69-72 



72 



THE CESTODES OF SEALS FROM THE ANTARCTIC 147 

and Diphyllobothrium by characteristic differences in the structure of the scolex, in 
the development of the muscles, and in the distribution of the testes. 

? Diphyllobothrium larvae 

[PL. 21, FIGS. 74-75] 

Host : Weddell seal (Leptonychotes weddelli) : Leopard seal (Hydrurga leptonyx) . 
Locality : Melchior Archipelago ; Debenham Islands ; Cooper Bay, South Georgia. 

Beside the adult forms enumerated in this paper, juvenile stages have also been 
discovered in five Weddell seals and in one Leopard seal. The infection bears in some 
cases a mass character. The identification of these juvenile forms was not possible, 
because of a complete lack of morphological features. They belong probably to one 
of the species dealt with here, most likely to Diphyllobothrium wilsoni or D. mobile. 

Based on differences in the scolex, two types of larvae might be distinguished : one 
with a kind of papillary modification occurring at the edge of the bothria, and another 
with the typical scolex of Diphyllobothrium but with the bothria smooth. 

The maximum length of the body of the first type, from the Weddell seal, is about 
3 mm. long and about 1 mm. in width, and the second, from the Leopard seal, is about 
i-2 mm. in length and 195 \x in width. 

? Diphyllobothrium sp. (larva) 

[PL. 21, FIG. 73] 

Host : Crabeater seal (Lobodon carcinophagus) . 
Locality: South Sandwich. 

This juvenile stage was collected by the Discovery on 21 March 1928 from the 
intestine of a Crabeater seal. 

The total length of the body is 7 mm. and the width 480 /jl. 

The scolex is well separated from the rest of the body and is heart-shaped, tapering 
anteriorly. Its length is 675 \l and its width 480 \i. 

The bothria are well developed. 

The unsegmented body has neither genital organs nor their rudiments, making 
identification impossible. 



OCCURRENCE OF THE PARASITE AND ITS RELATIONSHIP TO 

THE HOST 

As already stated, there are nine species of Pseudophyllidean Cestodes known to 
occur in the Antarctic seals. An analysis of the parasites and their hosts is shown in 
Table No. 4. 

Our knowledge of these parasites is very limited and the present material is merely 
the tenth collection of this kind. Nevertheless, certain speculations may be justifiably 
made on the occurrence and the host specificity of these parasites, as well as on their 



148 



THE CESTODES OF SEALS FROM THE ANTARCTIC 



Table No. 4 

Composition of Pseudophyllidean Cestodes in the present material 



Diphyllobothrium lashleyi (Leiper & Atkinson, 1914 
D. mobile (Rennie & Reid, 191 2) . 
D. quadratum (Linstow, 1892) 
D. scoticum (Rennie & Reid, 1912) 
? D. sp. (larva) 
D. wilsoni (Shipley, 1907) 
Glandicephalus antarcticus (Baird, 1853) 
G. perfoliatus (Railliet & Henry, 191 2) 
Baylisia baylisi gen. nov., spec. nov. 
Baylisiella tecta (Linstow, 1892) gen. nov 



Leptonychotes weddelli 

Hydrurga leptonyx 

»» »« 

Lobodon carcinophagus 
Leptonychotes weddelli, Hydrurga leptonyx 
Ommatophoca rossi 
Leptonychotes weddelli 
Lobodon carcinophagus 
Macrorhinus leoninus 



host-relationship. It may be presumed that Glandicephalus perfoliatus and Diphyl- 
lobothrium lashleyi are specific to the Weddell seal, G. antarcticus to the Ross seal, 
and D. quadratum and D. scoticum 1 to the Leopard seal. The newly-described Baylisia 
baylisi is the first identified Cestode from the Crabeater seal, although unidentified 
tapeworms have been reported from this host by Railliet & Henry (1912). Baylisiella 
tecta seems to be closely associated with the Elephant seal. Linstow (in Shipley, 1902) 
identified some Cestodes collected from the Ross seal as belonging to this species, but 
this identification requires confirmation. The two next species, Diphyllobothrium 
mobile and D. wilsoni, are less selective in their hosts, the first occurring in the 
Weddell and Ross seals, the second in the Leopard and Weddell seals. According to 
information supplied by Dr. G. C. L. Bertram, who collected the Graham Land 
material, the most frequently and heavily infested species are the Weddell seal, the 
Leopard seal, and the Elephant seal. In the Crabeater seal infestation with tape- 
worms very seldom occurs and the present record is virtually the first. 

It is obvious that the nature of the food has an enormous influence on the kind 
and number of parasites. The Weddell seal eats fish and cephalopods. The food of 
the Leopard seal is composed of penguins and fish. The food of the Crabeater seal 
consists of Crustacea, mainly Euphausiids. 

These Cestodes occur in specific parts of the gut of the host. Glandicephalus 
perfoliatus infests mainly the bile-duct, overhanging into the lower part of the 
intestine. It occurs very often as a mass infection, choking the lumen of the bile-duct. 
Diphyllobothrium lashleyi, D. mobile, D. quadratum, D. scoticum, and D. wilsoni infest 
the duodenal part of the gut and Baylisia baylisi and Baylisiella tecta occur in the 
rectum. Except for these two and Diphyllobothrium scoticum, the rest of the species 
occur very often as a mass infestation occupying almost all the free surface of the gut. 

D. scoticum does not occur in such numbers as the others but makes up for it in 
the size of the individuals. It is the largest Cestode recorded from the Antarctic seals. 
Baylisiella tecta also is fairly large and is not numerous in the gut. 

Diphyllobothrium wilsoni and D. mobile have been considered by previous authors 
as dwarfs, and the smallest species of Diphyllobothrium, D. wilsoni, is very small, 
reaching in mass infestation no more than 10 mm. in length. On the other hand, 
specimens collected from the less heavily infested Leopard seals reach from 5 to 9 cm. 

1 See footnote on p 137. 



PLATE 21, FIGS. 73-75 

Fig. 73. ? Diphyllobothrium sp. from Crabeater seal. 
Figs. 74-75. Two types of larvae from Weddell seal. 




PLATE 21 





73 



74 



Figs. 73-75 



THE CESTODES OF SEALS FROM THE ANTARCTIC 149 

in length. The larger specimens were considered by Shipley (1907) to be a separate 
species, D. scotti. It is quite probable that further investigation of the Cestodes of 
Antarctic seals will prove that the small size of D. mobile is also caused by the living 
conditions in mass-infected hosts. 

Where a mass infestation occurs one species of Cestode only is invariably present. 
Usually in the moderately infested hosts the Cestodes may represent more than one 
species. Glandicephalus perfoliatus was recorded in the same host together with 
Diphyllobothrium lashleyi, D. mobile, and D. wilsoni. The same has been shown for 
the occurrence of D. scoticum with D. quadratum and D. wilsoni in the same gut. 

No pathological changes of the gut have been observed, except in the rectum of 
the Elephant seal, where Baylisiella tecta provokes large nodules, about 3 cm. in 
diameter. This is caused by the scolex being very deeply embedded in the intestinal 
wall. Judging from the mass occurrence of parasites in the gut of seals, there must 
be a high degree of immunity on the part of the host against toxic factors provoked 
by the parasite. 

It seems from the literature that the Pseudophyllidean Cestodes found in the 
Antarctic seals do not occur in any other species of Pinnipeds. 

REFERENCES 

Ariola, V. 1900. Revisione della famiglia Bothriocephalidae s. str. Arch. Parasit. Paris, 3: 

369-484. 
Baird, W. 1853a. Descriptions of some new species of Entozoa from the collection of the British 

Museum. Proc. zool. Soc. Lond. 21: 18-25. 
1853. Catalogue of the species of Entozoa or intestinal worms contained in the collection of the 

British Museum. 132 pp. London (British Museum (Nat. Hist.)). 
1855. Catalogue of the species of Entozoa or intestinal worms contained in the collection of 

the British Museum. Ann. Mag. nat. Hist. (2) 15: 69-76. 
Baylis, H. A., in Hamilton, J. E. 1934. The southern sea lion Otaria byronia (De Blainville). 

'Discovery' Rep. 8: 306. 
Bertram, G. C. L. 1940. The biology of the Weddell and Crabeater seals. British Graham Land 

Expedition, 1934-1937. Sci. Rep. 1. (1) : 1-139. 
Blanchard, R. 1894. Notices sur les parasites de l'homme, 3 me serie; Sur Krabbea grandis et 

remarques sur la classification des Bothriocephalines. C.R. Soc. Biol. Paris, 46: 699-702. 
Braun, M. 1897. Vermes. Bronn's Klassen, 4 Abt. lb: 1407-1454. 

1898. Vermes. Bronn's Klassen, 4 Abt. lb: 1535-1614. 

Cooper, A. R. 1921. Trematoda and Cestoda. Rep. Canad. Arct. Exped. 1913-1918. 9 (G-H) : 

1-28. 
Diesing, K. M. 1863. Revision der Cephalocotyleen. Abtheilung: Paramacocotyleen. S.B. 

Akad. Wiss. Wien 48, Abt. 1: 200-345. 

1856. Zwanzig Arten von Cephalocotyleen. Denkschr. Akad. Wiss. Wien 12: 23-38. 

Drummond, F. H. 1937. Reports of the expedition of the McCoy society for field investigations 

and research Lady Julia Percy Island. Proc. roy. Soc. Vict. 49: 401-406. 
Fuhrmann, O. 1920. Die Cestoden der Deutschen Sudpolar-Expedition 1901-1903. Deutsche 

Sudpol. -Exped. igoi-igo3 (Drygalski), 16: 469-524. 

1931. Cestoidea. Handb. Zool. Berl. 2: 416 pp. 

Germanos, N. K. 1895. Bothriocephalus schistochilos n. sp., ein neuer Cestode aus dem Darm von 

Phoca barbata. fena. Z. Naturw. 30: 1-38. 
Hamilton, J. E. 1934. The Southern sea lion Otaria byronia (De Blainville). 'Discovery' Rep. 

8: 269-318. 



150 THE CESTODES OF SEALS FROM THE ANTARCTIC 

Johnston, T. H. 1937. Entozoa from the Australian hair seal. Proc. Linn. Soc. N.S.W. 62: 

9-16. 
1937. The Cestoda of the Australasian Antarctic Expedition. Sci. Rep. Aust. Antarctic 

Exped. 10 (4) : 1-74. 
Joyeux, Ch., & Baer, J. G. 1936. Cestodes. Faune Fr. 30: 613 pp. 
Leiper, R. T., & Atkinson, E. L., 19 14. Helminthes of the British Antarctic Expedition 1910- 

191 3. Proc. zool. Soc. Lond. 1914: 222-226. 
1915. Parasitic worms with a note on free-living Nematode. Brit. Antarctic (' Terra 

Nova') Exped., 1910 Nat. Hist. Rep. Zool. 2 (3): 19-60. 
Leon, N. 1910. Un nouveau cas de Diplogonoporus brauni. Zbl. Bakt. 55: 23-27. 
Linstow, O. 1878. Compendium der Helminthologie. xxii + 382 pp. Hannover. 
1892. Helminthen von Siid-Georgien. Nach der Ausbeute der deutschen Stationen von 

1882-1883. Jb. Hamburg. Wiss. Anst. 9: 59~77- 
in Shipley, A. E. 1902. Nematoda, Cestoda. Rep. Coll. Nat. Hist. 'Southern Cross'. 

ix+344 PP- London. 
Luehe, M. 1899. Zur Anatomie und Systematik der Bothriocephaliden. Verh. dtsch. zool. Ges. 

1899: 30-55- 
Markowski, S. 1949. On the species of Diphyllobothrium occurring in birds, and their relation 

to man and other hosts. /. Helminth. 23: 107-126. 
Matz, F. 1892. Beitrage zur Kentniss der Bothriocephalen. Arch. Naturgesch. 1: 97-122. 
Meggitt, F. J. 1924. The Cestodes of Mammals. 282 pp. London. 
1924. On the life history of a reptilian tapeworm (Sparganum reptans). Ann. trop. Med. 

Parasit. 18: 195-204. 
Moniez, R. 1896. Traite de parasitologic animate et vegetate applique a la medecine. 680 pp. Paris. 
Mueller, J. F. 1937. A repartition of the genus Diphyllobothrium. J. Parasit. 23: 308-310. 
Nybelin, O., in Skottsberg: 1931. Saugetier- und Vogelcestoden von Juan Fernandez. Nat. 

Hist. Juan Fernandez and Easter Island. 3: 493-524. 
Railliet, A., & Henry, A. 1912. Helminthes recueillis par l'Expedition Antarctique francaise 

du ' Pourquoi-Pas ? ' II. Cestodes des phoques. Bull. Mus. Hist. nat. Paris. 18: 153-159. 
Rennie, J., & Reid, A. 1912. The Cestodes of the Scottish Antarctic Expedition (Scotia). 

Trans, roy. Soc. Edinb. 48: 441-453, and in Rep. Scient. Res. Voyage 'Scotia' (1902-1904), 

6: 243-256. 
Shipley, A. E. 1905. Notes on collection of Parasites belonging to the Museum of University 

College, Dundee. Nematoda, Cestoda, with hosts. Proc .Camb. phil. Soc. (biol.) 13: 95-102. 
1907. 'Cestoda.' Nat. Antarct. Exped., 1901-1904. Nat. Hist. 3: 6 pp. (British Museum 

(Nat. Hist.)). 
Southwell, T., & Walker, A. J. 1936. Notes on a larval Cestode from a Fur-seal. Ann. trop. 

Med. Parasit. 30: 91-100. 
Stiles, C. W., & Hassall, A. 1912. Index-catalogue of medical and veterinary Zoology: 

Cestoda and Cestodaria. Bull. U.S. hyg. Lab. 85: 1-467. 
Stunkard, H. W. 1947. On certain Pseudophyllidean Cestodes from Alaskan Pinnipeds. /. 

Parasit. 33: (suppl.) : 19. 

1948. Pseudophyllidean Cestodes from Alaskan Pinnipeds. /. Parasit. 34: 211-228. 

1949. Diphyllobothrium stemmacephalum Cobbold, 1858 and D. latum (Linn., 1758). 

/. Parasit. 35: 613-624. 
& Schoenborn, H. W. 1936. Notes on the structure, distribution and synonymy of 

Diphyllobothrium lanceolatum. Amer. Mus. Novit. 880: 1-9. 
Wardle, E. R. A., McLeod, J. A., & Stewart, I. E. 1947. Liihe's ' Diphyllobothrium ' (Cestoda) . 

/. Parasit. 33: 319-330. 
Zschokke, F. 1903. Die arktischen Cestoden. Fauna arct., Jena 3: 1-32. 




PRESENTED 

2 JUL 1952 



PRINTED IN 

GREAT BRITAIN 

AT THE 

UNIVERSITY PRESS 

OXFORD 

BY 

CHARLES BATEY 

PRINTER 

TO THE 

UNIVERSITY 



i 



THE 'MANIHINE' 
EXPEDITION TO THE 
GULF OF AQABA ^ 
1948-1949 




BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ZOOLOGY Vol. 1 No. 8 

LONDON : 1952 



THE 'MANIHINE' EXPEDITION 

O 

TO THE GULF OF AQABA 1948-1949 



CONTENTS 

I. Foreword: Station List and Collectors' Notes 
II. Preliminary Hydrological Report: G. e. r. deacon 

III. Sponges: m. burton 

IV. Turbellaria: Polycladida: s. prudhoe 
V. Gephyrea: a. c. Stephen 

VI. Mollusca: w. j. rees and a. stuckey 
VII. Echinodermata: a. m. clark 
VIII. Tunicata: w. g. van name 

IX. Fishes: n. b. Marshall 

Pp. 151-252, Pis. 22-32; 10 Text-figs 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ZOOLOGY Vol. I, No. 8 

LONDON: 1952 



THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted in 1949, is issued 
in five series, corresponding to the Departments of the 
Museum. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four hundred 
pages, and will not necessarily be completed within one 
calendar year. 

This paper is Vol. 1, No 8, of the Zoology series. 










PRINTED BY ORDER OF THE TRUSTEES OF 
THE BRITISH MUSEUM 

Issued August 1952 Price Twenty-five shillings 



THE 'MANIHINE' EXPEDITION TO THE 
GULF OF AQABA 

I. FOREWORD: STATION LIST AND 
COLLECTORS' NOTES 

During the winter of 1948-1949 the motor-yacht Manihine was engaged in biological 
investigations in the Gulf of Aqaba on behalf of the British Museum (Natural 
History), the work being under the supervision of Mr. N. B. Marshall. 

This gulf is of special interest because in it the peculiarities of the Red Sea appear 
at their most intense. The Red Sea is geologically young with a fauna derived from 
that of the Arabian Sea and, possibly, the Mediterranean. This immigrant fauna is 
now completely isolated from the last-mentioned and also partially isolated from the 
former by reason of the narrowness and shallowness of the connecting passage, 
the Strait of Bab-el Mandeb. It also finds itself in a region where some, at least, of 
the ecological conditions are very different. The most noticeable of these ecological 
differences is to be found in the isohaline and isothermal nature of the water below 
200 metres and the complete absence of any cold, deep-water layer. The John Murray 
Expedition (Seymour Sewell, 1935, John Murray Exp., Reports, 1, 1) recorded tem- 
peratures from 21-64 to 21-84° C. at depths of 1,000 to 1,900 metres in the Red Sea, 
but at similar depths in the Gulf of Aden the temperature was at least io° C. lower 
(3*59 _II, 53° C.)- The degree of isolation of the Aqaba fauna is greater than that 
of any other part of the Red Sea since the passage between the two, the Strait of 
Tiran, provides only a restricted channel for faunal interchange. The strait is both 
narrow and shallow, forming a distinct sill, with a greatest depth of less than about 
300 metres ; on either side of the sill the water deepens rapidly to 1,000 metres and 
upwards. The hydrological conditions inside the gulf appear to be essentially similar 
to those in the Red Sea proper, though, as might be expected, salinities are some- 
what higher. 

In this Bulletin are reports on some of the collections that were brought back. 
Other reports, including a study of the interchange of heat and water vapour between 
the surface water and the air, will be prepared as opportunities offer, but in some 
instances the collections will be studied in conjunction with other material and will 
not form the subject of a special report. 

Acknowledgements and thanks are due to many individuals and institutions whose 
material aid or advice contributed greatly to the expedition. Foremost among them 
is Major H. W. Hall, O.B.E., M.C., who not only provided the ship and was respon- 
sible for most of the preliminary organization, but who, with Mrs. Hall, accompanied 
the expedition taking a large share of the actual collecting and doing most of the 
photography. A small selection of the photographs is published here to give a general 
impression of the gulf and its surroundings. Many localities could not have been 
visited but for the skilful pilotage of Captain Hargreaves through poorly charted 



154 THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 

waters, and to him, and to his hard- working crew, all possible thanks are due. The 
Hydrographer of the Navy and the Director of 'Discovery Investigations' lent 
apparatus vital to the expedition and His Excellency the Egyptian Ambassador in 
London made arrangements that ensured pleasant and harmonious relations wherever 
the ship was in Egyptian waters. Lastly, thanks are due to the Government Chemist, 
whose department carried out the analyses of salinities. 

Except for the plankton and some of the fishes all material was obtained from 
littoral areas and coral reefs (or coral patches). Localities where collections were 
made are indicated on the chart. Within the Gulf of Aqaba (reading from north 
to south) these were : 

Aqaba (PL 22, fig. 1) Hobeik (PI. 23, fig. 4) 

Faraun Island (PI. 22, fig. 2) Dahab (PI. 24, fig. 5) 

Graa Um Nageila (PI. 24, fig. 6) 

Mualla (PI. 23, fig. 3) Abu Zabad 

Along the Sinai shores there are well-formed coral reefs at Dahab and from Um 
Nageila southwards. The bulk of the invertebrate material was obtained from these 
regions, particularly from Abu Zabad on the 10th and nth February 1949 when 
there were low spring tides. North of Dahab there were coral patches at all localities 
visited, but these never become massed to form a definite reef. 

Outside the gulf collections were made at the following localities : 

Sanafir Island (PI. 25, fig. 7) Sherm-el-Moiya (PI. 26, fig. 9) 

Tiran Island (PI. 25, fig. 8) Ras Muhammad Bay (PI. 26, fig. 10) 

Sherm Sheikh 

Most time was spent on Sanafir Island, where there were well-formed coral reefc. 
Here, as elsewhere, much material was collected by diving for pieces of coral and 
extracting the small invertebrates and fishes. 

It will be observed that in the Station List no temperatures are given for depths 
below 40 metres. It was, however, established that at all stations where deep-water 
samples were taken (i.e. where salinity figures are given in the list) the temperature 
exceeded 18-5° C. 

The following are the meanings of the abbreviations used in the list. 

D.M. = Dredge, medium. 

D.S. = Dredge, small. 

K.T. = Kelvin tube. 

N. 70 V. = Vertical haul by silk plankton net with mouth 70 cm. diameter. 

N. 100 V. = Vertical haul with stramin plankton net with mouth 100 cm. 

diameter. 
O.T.L. = Otter trawl, large. 







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Legends to Plates 22-2J. 



PLATE 22 

Fig. i. Aqaba looking north-east. 

Fig. 2. Gezeret-el-Faraun from the south-east. 

PLATE 23 

Fig. 3. Looking north from the anchorage at Mualla. 
Fig. 4. Hobeik. 

PLATE 24 

Fig. 5. Typical gulf scenery. Coast 5 miles south of Dahab. 
Fig. 6. Mangrove swamps at Um Nageila. 

PLATE 25 

Fig. 7. Sanafir Island ; Fish-eagle's nest. 
Fig. 8. Tiran Island, seen from Sanafir. 

PLATE 26 

Fig. 9. Sherm-el-Moiya ; looking north-east from the entrance. 
Fig. 10. Manihine at anchor in Ghazulani Bay with Ras Muhammad in 

the distance. 

PLATE 27 

Fig. 11. Abandoned police post at Naweibi-el-Terabin, about 45 miles 

south of Aqaba. 
Fig. 12. Arab fisherman using cast net. 



Bull. B.M. (N.H.) Zoology, I, 8 



PLATE Tl 




Fig. i. AQABA 







Fig. 2. GEZERET-EL-FARAUN 



Bull. B.M. {N.H.) Zoology, I, 8 



PLATE 23 




Fig. 3. MUALLA 




Fig. 4. HOBEIK 



Bull. B.M. {N.H.) Zoology, I, 8 



PLATE 24 




Fig. 5. GULF SCENERY NEAR DAHAB 




Fig. 6. UM NAGEILA 



Bull. B.M. (N.H.) Zoology, 1, 8 



PLATE 25 



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Fig. 8. TIRAN ISLAND 



Bull. B.M. (N.H.) Zoology, I, 8 



PLATE 26 




Fig. 9. SHERM-EL-MOI YA 




Fig. io. GHAZULANI BAY 



Bull. B.M. {N.H.) Zoology, I, 8 



PLATE 27 




Fig. ii. NAWEIBI-EL-TERABIN 



&*L*rl*±. • ' 




Fig. 12. CAST NET 



II. PRELIMINARY HYDROLOGICAL REPORT 

By G. E. R. DEACON, F.R.S. 

NATIONAL INSTITUTE OF OCEANOGRAPHY 

The observations confirm the general picture of the water circulation described by 
A. F. Mohammed in Proc. Roy. Soc. B. 128, 1939, and give some new information 
about the surface layer. 

As plotted in Fig. 1, the surface water at Stations 30 and 31 in the Straits of 
Tiran, and at Station 23 twenty miles farther north on the east side of the gulf, had a 
salinity less than 40-6% which can be attributed to the inflow of water from the Red 
Sea. There is some indication that the inward movement has a greater influence on 
the east side of the gulf since the surface salinity at Station 17 nearly half-way up the 
gulf is only 40-63% . For the remainder of the gulf, including all stations north and 
west of a line from Station 26 to Station 17, the water between the surface and 
20 fathoms can be regarded as almost isothermal and isohaline, with a temperature 
of 21 to 22 C. (in January) and a salinity of 407 to 4o-8% . 

Excepting Stations 31, 30, and 25, the observed surface temperature appears to 
depend more on the time of day at which the measurement was made than the 
position of the station in the gulf. When plotted against time of day (Fig. 2) the 
temperatures lie fairly closely about a curve of diurnal temperature change which 
has a maximum at approximately 13.00 hours. The bathythermograph observations 
made at all the stations always show a temperature less than that measured by taking 
a surface sample and using a thermometer. Some of the differences can be attributed 
to the shallower depth of the sample scooped up in a surface sampler, and to the 
existence of an appreciable thermal gradient in the first foot or two of water. The 
differences between the thermometer and bathythermograph readings when plotted 
against the time of day (Fig. 3) lie fairly closely about a curve with a maximum of 
o*55° C, which is very similar to that showing the diurnal temperature variation 
(Fig. 2) at 13.00 hours. The differences between the readings at the surface and a 
depth of 40 metres on the bathythermograph slides (Fig. 4) shows that this differ- 
ence, which varies between 0-2° and o-6° C, varies according to a similar curve. 

It is expected that some further information about the interchange of heat and 
water vapour between the surface water and the air can be obtained from the data, 
and, when some attempt is made to smooth out the diurnal temperature variations, 
one or two useful indications of the surface movements ; but the best that can be 
done at present is to regard the upper 40 metres of water as more or less uniform, 
excepting Stations 31, 30, and 23. These appear to be influenced by the inflow of 
surface water from the Red Sea. Reference to Fig. 1 will also show that the stations 
near the eastern shore in the southern part of the gulf appear to be influenced by a 
more recent inflow of water than those farther north and west. 



zoo. 1. 8. 




Fig. i. Surface salinities. The underlined figures are the 
station numbers. 



THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 



161 



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Fig. 2. Surface temperature in relation to time of day. (Numbers refer to stations.) 



1 62 



THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 

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Fig. 4. Difference between temperature at surface and at 40 metres, plotted against time 

of day. (Numbers refer to stations.) 



III. SPONGES 

By MAURICE BURTON, D.SC. 

The sponges represent thirty-three species, and although their study has resulted 
in little of unusual interest, a useful addition to the faunal list of the Red Sea area 
has been made. In addition, it has been possible to establish the correct identity 
of some of the forms described by Keller (1889 and 1891), which has long been in 
doubt. Most of the thirty-three species are common to the Indian Ocean fauna, some 
having been recorded also from Australia or the Indo-Pacific. It is of interest to note, 
however, that twelve species appear to be endemic, but this may be due largely to 
gaps in our knowledge of the Indian Ocean fauna. Furthermore, there are three 
species (Leuconia nausicae, Tethya aurantium, and Pseudosuberites mollis) belonging 
more properly to the Mediterranean fauna. 

The commonest form in the Gulf of Aqaba seems to be Callyspongia viridis, which, 
according to the members of the expedition, is 'abundant everywhere'. 

LIST OF SPECIES AND SYSTEMATIC NOTES 
Order CALCAREA 

Leucosolenia canariensis (Michlucho-Maclay) 

Nardoa canariensis Michlucho-Maclay, 1868: 221. 
Leucosolenia canariensis, Dendy & Row, 1913: 724. 

Occurrence. Mualla, 30x49, under rocks at low tide ; Sherm-el-Moiya, 3.U.49. 
Remarks. A greyish white, typical specimen, 10 mm. across. 
Distribution. Arctic; Mediterranean; Cape Verde Islands; Canaries; Red Sea; 
Mauritius ; NW. Pacific (Commandorski Islands) . 

Leucosolenia tenuipilosa Dendy 

Leucosolenia (Clathrina) tenuipilosa Dendy, 1905 : 227, pi. xiii, fig. 9. 
L. canariensis (pars), Thacker, 1908: 762. 
Clathrina tenuipilosa, Row, 1909: 185. 
Leucosolenia tenuipilosa, Dendy & Row, 1913: 723. 

Occurrence. Dahab, 14.ii.49 ; Abu Zabad, 11.ii.49. 

Remarks. There are a number of typically cushion-shaped specimens, up to 30 mm. 
across, which were brown or fawn in formalin, and now, in spirit, are coloured a 
greyish brown. 

Distribution. Ceylon; Red Sea; Cape Verde Islands. 

Gr antes sa glabra Row 

Grantessa glabra Row, 1909: 203, pi. xix, figs. 5-6; Dendy & Row, 1913: 752. 

Occurrence. Sherm Sheik, n.i.49; Abu Zabad, 10.ii.49, on ree f a * l° w tide. 
Distribution. Red Sea. 



164 THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 

Leuconia bathybia (Haeckel) 

Dyssycum bathybia Haeckel, 1869: 241. 

Leucaltis bathybia, idem, 1872: 156, pi. xxviii, fig. 2. 

Leucandra bathybia, Dendy & Row, 191 3: 773. 

Occurrence. Sherm Sheik, 2.ii.49, 2 fms. ; Sanafir, 6.U.49. 

Remarks. The four specimens may possibly represent two well-marked varieties, 
and, since the species was originally subdivided in this manner, it may be worth 
while to consider them in this light. 

The first specimen is the smaller, a few millimetres high, and of typical form and 
colour. The skeleton is arranged as Haeckel described it, and the rays of the large 
quadriradiates have a maximum of 0-4 by 0-032 mm. 

The other three range from a few millimetres high to 16 mm. high by 12 mm. 
diameter. Again, the external form is typical, as well as the spiculation. But in 
these three the rays of the quadriradiates have a maximum of 0-96 mm. by 0-09 mm. 

Either the first of the present specimens represents Haeckel's var. perimina and 
the other three var. arabica, or, what is much more likely, we have to deal with a 
species showing a tendency to vary widely in the measurements of the spicules. 

The first specimen and two out of the group of three were found at the same 
station, Sherm Sheik. 

Distribution. Red Sea ; ? Australia. 

Leuconia nausicae (Schuffner) 

Leucaltis nausicae Schuffner, 1877: 407, pi. xxiv, fig. 1. 
Leucandra nausicae Dendy & Row, 1913: 774. 

Occurrence. Sanafir, 9.L49 ; Tiran, io.i.49 ; Abu Zabad, 11.ii.49, on reef at low tide. 

Remarks. The two specimens seem to agree closely with the description of the 
holotype, which is the only other recorded specimen. Presumably Row (I.e.) examined 
this and, as a consequence, the species was transferred to Leucandra. It is difficult, 
therefore, to accept Topsent's (1937: 14) remark that 'Leucaltis Nausicae Schuffner 
se confond vraisemblablement avec Leucetta solida (O. Schmidt) '. 

Distribution. Mediterranean. 

Kebira uteoides Row 

Kebira uteoides Row, 1909: 210, pi. xx, figs. 8-9, text-figs. 7-8; Dendy & Row, 1913: 785. 

Occurrence. Sherm Sheik, 2 fms., 2.U.49. 

Remarks. The single specimen, 20 mm. high, is typical, in both external appear- 
ance and the details of the skeleton. 

Distribution. Red Sea. 

Order TETRAXONIDA 

Stelletta purpurea Ridley 

(For synonymy see Burton, 1926.) 

Occurrence. Tiran, io.i.49 > Sanafir, 8 and 9.L49 and 4.H.49 ; Sherm-el-Moiya, 3.U.49. 
Remarks. The spiculation of the several specimens shows the usual variation in 



THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 165 

size. The main interest lies, however, in the external form. The smallest specimens, 
10 to 15 mm. diameter, have the spherical or subspherical shape typical of the species, 
but in one or two cases these small spherical sponges have coalesced to give an ir- 
regular lobulated mass. In the larger specimens, 50 to 60 mm. across, on the other 
hand, the form is often extremely irregular, suggesting not only the coalescence of 
several smaller sponges but irregularities of growth due to environmental factors. 
Distribution. Red Sea ; Indian Ocean ; Malay ; Australasia ; Antarctic. 

Chondrilla sacciformis Carter 
(For synonymy see Burton, 1924.) 

Occurrence. Sherm-el-Moiya, 3.^.49. 
Distribution. Indian Ocean ; Malay. 

Chondrosia reniformis Nardo 
Chondrosia reniformis Nardo, 1847: 272. 

Occurrence. Abu Zabad, n.ii.49. 

Remarks. The two specimens appear to be typical except that there is a sparse 
accumulation of fine sand grains in the outer layers of the cortex. 

Distribution. Atlantic coast of Europe ; Mediterranean ; South Africa (Stil Bay) ; 
Indian Ocean ; Malay ; Australia. 

Chrotella cavernosa (Lamarck) 

Tethya cavernosa Lamarck, 1813: 70; 1815: 383. 
T. cranium var. australiensis Carter, 1886: 127. 
Cinachyra australiensis, Burton, 1934: 523. 
(For further synonymy see Burton, I.e.) 

Occurrence. Mualla, 30^.49, at low tide under rocks. 

Distribution. Red Sea ; Indian Ocean ; Malay ; Australia ; Philippines. 

Tethya aurantium (Pallas) 

(See Burton 1924 and 1949: 122.) 

Occurrence. Sherm Sheik, 2.U.49, n.i.49, and 2.U.49 ; Tiran, io.i.49; Mualla, 
30^.49, at low tide under rocks. 

Remarks. The five specimens, all somewhat flattened, are fawn, orange, or red 
(in formalin) and measure 7, 8, 12, 18, and 21 mm. across respectively. 

Distribution. Arctic ; North Atlantic ; West Indies ; Mediterranean. 

Tethya robusta Bowerbank 
(For synonymy see Burton, 1924.) 

Occurrence. Mualla, 30^.49, under rocks at low tide; Abu Zabad, 10 and 11.ii.49, 
on reef at low tide. 

Remarks. The six specimens measure 13, 15, 21, 25, 26, and 28 mm. across respec- 
tively. The colour (in formalin) is pink to red. There is, however, another specimen 



166 THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 

consisting of five lobes set in a horizontal plane, each lobe being about 20 mm. across. 
Its colour was a cerise-red in formalin. Clearly this specimen has been formed by the 
complete coalescence of five adjacent individuals. It is not unknown for two speci- 
mens to fuse in this way, but five is unusual. 

The spiculation is typical in all but two specimens, which lack the larger micrasters. 
In other words, these two should be assigned to Tethya japonica Sollas. In 1924 I 
suggested that this so-called species was probably a reduced form of T. diploderma 
Schmidt (= T. ingalli Bowerbank), but it now seems that it is a mixture of the 
reduced forms of both T. robusta and T. ingalli. 

Distribution. Australia ; Malay ; Indian Ocean. 

Pseudosuberites mollis Topsent 

Pseudosuberites mollis Topsent, 1925 : 9, fig. 2m. 

Occurrence. Mualla, 30.1.49, under rocks at low tide. 

Remarks. The sample consists of three fragments of a soft and delicate sponge, 
having approximately the characters described by Topsent (I.e.). The spicules are 
slightly larger, 0-15 to 0-45 by 0-005 to 0-008 mm., as compared with 0-175 to 0-315 
by 0-0065 mm - m * ne holotype, but the variations in the shape of the spicules are 
similar to those figured by Topsent. 

Distribution. Mediterranean (Etang de Thau). 

Haliclona toxophorus (Hentschel) 

Gellius toxophorus Hentschel, 1912: 392, pi. xxi, fig. 46. 
G. toxotes, idem, I.e.: 392, pi. xxi, fig. 47. 

Occurrence. Sherm Sheik, n.i.49. 

Remarks. The two small fragments are evidently from one sponge which formed a 
flattened, massive incrustation, with oscules slightly raised. Almost transparent, 
soft and compressible, delicate in texture, the specimen appears to be denuded of 
flesh, the skeleton, an isodictyal and unispicular network, being held together by 
spongin at the nodes. The megascleres are oxea, with a tendency to become strongy- 
lote at one or both ends, 0-24 by 0-012 mm. The microscleres are toxa, 0-02 to o-i mm. 
across. 

The two species described by Hentschel were sufficiently closely related, judging 
by the original descriptions, to suggest their identity one with the other. The inter- 
mediate character of the present material adds point to this. 

Distribution. Malay. 

Adocia dendyi (Burton) 

Toxochalina robusta Dendy, 1905: 139; idem, 1921: 29. 
T. dendyi Burton, 1931: 340, fig. ib. 
Nee Toxochalina robusta Ridley. 

Occurrence. Sherm Sheik, n.i.49. 

Remarks. The several specimens are all small and cushion-shaped, with con- 
spicuous oscules 2 to 3 mm. diameter. The colour, in spirit, is brownish grey, and 



THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 167 

the texture soft, compressible, elastic. The main skeleton is a close-meshed reticula- 
tion of fibres, the ascending fibres multispicular (3 to 4 spicules), the connectives 
unispicular. The tangential dermal skeleton is very much as figured by me (I.e., 
fig. 2b) and is unispicular. The spicules are oxea o-i by 0-004 mm., and toxa of about 
the same length. 

Distribution. Indian Ocean. 

Callyspongia viridis (Keller) 
Dactylochalina viridis Keller, 1889: 391, pi. xxiii, figs. 37-43. 

Occurrence. Sherm Sheik, 2 and 3.H.49 ; Tiran, io.i.49 ; Abu Zabad, 10 and 11.ii.49, 
on reef at low tide; Dahab, i3.i.49 and 14.ii.49; Sanafir, 4, 5, and 6.U.49. 

Remarks. Of the eleven specimens, only one is almost identical with that figured 
by Keller (I.e., fig. 37), nine of the remainder being irregularly massive, on the whole 
smaller, and the eleventh being no more than a thin incrustation on a coral. All have 
the typical vents and the typical pore-sieves (Keller, I.e., fig. 40), although in some 
cases the pore-sieves are less strongly marked. In a few cases, at least, the characters 
of the surface have been blurred by preservation in formalin. 

The characters of the skeleton are comparatively uniform for the nine irregularly, 
massive specimens, but the typical specimen and the thin incrustation show features 
which merit special notice. In the nine specimens the network of the main skeleton 
consists of well-marked primary or ascending fibres which branch, as they run to the 
surface, in a somewhat irregular manner. At the centres of the fibres is a more or 
less continuous core of spicules arranged in an untidy manner (almost irregularly 
sub-plumose), often with individual spicules projecting from the fibres. The primary 
fibres are connected by secondary fibres, thinner than the primaries, and forming 
often an irregular network. In these the spicules are arranged, usually, uniserially ; 
but, again, individual spicules may project, at right angles to the main series, beyond 
the surface of the fibres. The tangential skeleton at the surface is a close-meshed 
network of fibres, cored by uniserially arranged oxea, and showing no obvious 
differentiation into primary and secondary meshes. The average diameter of the 
meshes is 0-04 mm. The oxea vary from 0-08 to 0-16 by 0-004 t° 0-005 mm. 

The main skeleton of the one typical specimen (i.e. externally typical) is unlike 
that of the nine specimens in that it approaches the ceraochalinoid condition. 
It is a very close-meshed reticulation of thick fibres which appear at first sight to be 
aspiculous. In general it resembles that shown in Keller's fig. 39. On closer examina- 
tion, however, it can be seen that the spicules are present, are reduced in numbers, 
and seldom more than 0-002 mm. thick ; and often a spicule may be discontinuous 
throughout its length (as though breaking up) . 

As a result of comparing the external forms of these sponges, as well as the struc- 
ture of their skeletons, there seems little doubt that they are all conspecific and that 
the variation in their skeletons is unimportant. Generally speaking, it seems that in 
the younger sponges and the newer tissues the reticulation of the fibres is more loose 
and the fibres themselves more heavily cored with spicules ; that with maturity the 
skeleton is more closely knit and the proportion of spicule to spongin decreases (cf . 

zoo. 1. 8. z 



168 THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 

Burton, 1926 : 265) . One further point may be mentioned. In the specimen, described 
above as typical, the spicules have the appearance, as a result of their slender build 
and the discontinuous structure already referred to, of being dissolved or absorbed. 
Whether, in fact, this is the case is, however, problematical. 

The colour of the present specimens, in formalin, was grey to fawn. 

Distribution. Red Sea. 

Gelliodes fibulatus Ridley 

Gelliodes fibulatus Ridley, 1884: 427, pi. xxxix, fig. 1, pi. xli, fig. b; Ridley & Dendy, 1887: 47, 

pi. xii, fig. 2; Lendenfeld, 1887: 793. 
Pachychalina fragilis, Lindgren, 1897: 481; idem, 1898: 290. 
Gelloides ramosa Kieschnick, 1898: 47. 
? Pachychalina conulosa, idem, I.e.: 51. 
Gelliodes ramosa, idem, 1900: 565, pi. xliv, fig. 3. 
? Pachychalina conulosa, idem, I.e. : 568, pi. xliv, fig. 8. 
Gelliodes fibulatus, Hentschel, 19 12: 393. 
Sigmaxynissa fibulata, Burton, 1928: 115. 

Occurrence. Graa, 30.1.49; Sherm-el-Moiya, 3.^.49; Sanafir, 6.H.49. 

Remarks. It is somewhat surprising to find what appear to be typical examples of 
this species so far west as the Gulf of Aqaba. All records previously have been for 
the Malay region and the Indian Ocean (Andaman Islands). 

Distribution. Malay; Indian Ocean; (? Australia). 

Mycale euplectellioides Row 

Esperella euplectellioides Row, 191 1: 333, pi. xxxvii, fig. 12, text-fig. 16. 
Mycale euplectellioides, Burton, 1926: 80. 

Occurrence. Sherm Sheik, 2.H.49 ; Graa, 30.1.49; Dahab, 13.L49; Sanafir, 4 and 
6.U.49. 

Remarks. The sponge occurs in irregular masses on coral, the largest being some 
30 mm. across. Externally there is a close resemblance to the type, and from the 
condition of the several specimens, when removed from the formalin in which they 
were originally preserved, it is clear that a copious amount of mucus is present in life. 

The skeleton is typical except that microscleres are extremely rare, none being 
found except in a section from one specimen, which contained a few sigmata, 0-05 to 
0-08 mm. chord, and one anisochela 0-024 mm - chord. 

Distribution. Red Sea ; Suez Canal. 

Mycale (Carmia) suezza (Row) 

Esperella suezza Row, 191 1: 338, fig. 18. 

Occurrence. Mualla, 31.1.49; Dahab, 14.ii.49. 

Remarks. Two samples are assigned doubtfully to this species. The first is a thin 
incrustation, orange-coloured in formalin, and a larger, irregularly massive sponge, 
having the same colour and general appearance. The skeleton has the same structure 
as the holotype of Mycale suezza, but in neither specimen has it been possible to find 
a single microsclere. 

Distribution. Red Sea. 



THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 169 

Mycale (Aegagropila) erythraena (Row) 

Esperella erythraena Row, 191 1: 340, fig. 19. 
Mycale erythraena, Burton, 1926: 80. 

Occurrence. Dahab, 4.11.49. 

Remarks. The single specimen forms a thin, irregular incrustation on coral. Its 
colour, in formalin, was grey. The arrangement of the skeleton approximate closely 
to the type, and the megascleres are typical in form and size ; but in spite of repeated 
searching not a single microsclere has been found. 

Distribution. Red Sea ; Suez Canal. 

Genus PARISOCIELLA gen. n. 

Type Species. Esperiopsis anomala, Ridley & Dendy, 1886 : 341. 

Diagnosis. Mycaleae with skeleton an irregular reticulation of spongin fibres cored 
by slender tylostyli; microscleres, when present, degenerate anisochelae palmatae 
and toxa. 

Parisociella anomala (Ridley & Dendy) 

Esperiopsis anomala Ridley & Dendy, 1886: 341 ; idem, 1887: 84. 

Ceraochalina gibbosa Keller, 1889: 386, pi. xxiv, fig. 44. 

Ophlitaspongia arbuscula Row, 1911: 347, pi. xxxix, fig. 22, pi. xl, fig. 25, text-fig. 22. 

O. horrida, idem, I.e.: 349, pi. xl, fig. 26, text-fig. 23. 

Occurrence. Sanafir, 4 and 9.U.49, along the shore among rocks; Abu Zabad, 
10.ii.49, on reef at low tide. 

Diagnosis. Sponge typically branching, surface uneven, minutely hispid ; oscules 
not apparent ; texture soft, elastic ; colour alive red, in spirit greyish yellow to dark 
grey; main skeleton an irregularly isodictyal reticulation of fibres cored by mega- 
scleres ; dermal skeleton of radiating brushes of megascleres ; megascleres tylostyli, 
slender and often appearing as styli, 0-25 to 0-3 by 0-002 to 0-005 mm - 1 microscleres 
usually absent and never plentiful, anisochelae palmatae, o-oi mm. chord, and 
toxa, 0-02 to 0-06 mm. long. 

Remarks. The diagnostic features of this species are unsatisfactory, since the 
microscleres, even when present, exist in such small quantities and are difficult to 
find. Further, the main skeleton is so like that of Mycale euplectellioides, growing in 
the same habitat, that only the external form remains as a guide to identification. 
If, therefore, the particular specimen is macerated or fragmentary the possibility 
of wrong identification is great. 

The present three specimens include a fragment of a branch, which is macerated, 
and two extensive, but low, incrustations on pieces of coral. The colour, in formalin, 
was orange and yellowish brown, in spirit, yellow or brown. No microscleres were 
found. 

Distribution. Red Sea ; Honolulu. 

Lissodendoryx cratera (Row) 
Myxilla cratera Row, 191 1: 343, pi. xxxvii, fig. 13, text-fig. 20. 
Occurrence. Abu Zabad, 11.ii.49. 
Distribution. Red Sea. 



170 THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 

Agelas mauritianus (Carter) 

Ectyon mauritianus Carter, 1883: 310, pi. xii, fig. 3. 

Agelas mauritianus, Ridley & Dendy, 1887: 164, pi. xxix, fig. 10. 

A. cavernosa Thiele, 1903: 963, fig. 28. 

A. mauritiana, Dendy, 1905: 174. 

Occurrence. Sanafir, 6.11.49. 

Remarks. A fairly large fragment which, in formalin, was pink outside and orange 
in the interior. 

Distribution. Indian Ocean ; Malay. 

Halichondria glabrata Keller 
Halichondria glabrata Keller, 1891: 311, pi. xvi, fig. 9; Burton, 1926: 75. 

Occurrence. Abu Zabad, 11.ii.49. 

Remarks. A single, thinly encrusting specimen, in colour pale brown, both in 
formalin and in spirit. 

Distribution. Red Sea. 

Rhaphoxya typica Hallmann 

Rhaphoxya typica Hallmann, 1917: 643, pi. xxix, fig. 3, pi. xxxviii, figs. 8-9, pi. xxxix, fig. 5, 
pi. xlii, figs. 1-2, text-fig. 17. 

Occurrence. Sanafir, 6.U.49 ; Abu Zabad, 10.ii.49, on reef at low tide. 

Remarks. The several species which may be assigned to Rhaphoxya are mainly 
Australian and none has been previously recorded from the Red Sea, although 
Anacanthaea nivea Row might conceivably belong to this genus. Yet the present 
two specimens clearly belong to Rhaphoxya and are almost certainly conspecific with 
the genotype. They are both encrusting, but their general appearance and the 
characters of the surface agree closely with those described and figured by Hallmann, 
except that the pore-areas (?), in his pi. xxxviii, fig. 8, are not so numerous in the 
'Manihine' sponges. There is, also, a close agreement in the shape of the spicules 
and their arrangement in the skeleton, except that the trichites are not numerous 
and, as far as can be seen, do not form dragmata. 

A striking feature of the anatomy concerns the presence of numerous oval groups 
of cells, looking very like embryos, which they may well be, except that they vary 
somewhat in size, from 0-08 to 0-2 mm., with 0-12 mm. as the average, across the 
long axis. The tissues of the sponge contain numerous brown pigment cells in the 
surface layers, and the ' embryos ' lying in the surface tissues are also filled with them. 

Distribution. Australia. 

Order KERATOSA 

Aplysilla lacunosa Keller 
Aplysilla lacunosa Keller, 1889: 356, pi. xxii, figs. 19-22. 

Occurrence. Sanafir, 6.ii.49. 

Remarks. A single, very small, incrusting specimen, purple in colour, showing the 
typical fibres (see Keller, I.e., pi. xxii, fig. 22). 
Distribution. Red Sea. 



THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 171 

Megalopastas erectus Row 

Megalopastas erectus Row, 191 1: 360. 

Occurrence. Sherm Sheik, n.i.49; Dahab, 14.ii.49. 

Remarks. The two specimens form irregular encrustations, with the surfaces 
irregularly conulose. The colour of one, in formalin, was purple, in spirit it turned 
to a deep violet ; in the other it was fawn in formalin and the same in spirit. 

Distribution: Red Sea. 

Spongia officinalis Linnaeus, var. arabica (Keller) 
Euspongia officinalis, var. arabica Keller, 1889: 342; Topsent, 1906: 558; Row, 191 1: 379. 

Occurrence. Abu Zabad, 10 and 11.ii.49, on ree f a * low tide ; Sherm-el-Moiya, 3.ii.49 ; 
Sanafir, 9^.49. 

Remarks. There are two typical specimens, two very small specimens in which the 
skeleton only remains and which are doubtfully assigned to this species, a fifth, 
typical but very small, and a sixth specimen which agrees in general appearance, 
but has the internal tissues so crowded with sand that a better identification is not 
possible. 

The colour in formalin varies from fawn (the specimens without flesh) to dark 
brown. 

Distribution. Red Sea. 

Heteronema erecta Keller 

Heteronema erecta Keller, 1889: 340, pi. xx, figs. 4, 7, 8; Topsent, 1906: 558; Row, 191 1: 369. 
Duriella nigra Row, 191 1: 370, pi. xli, fig. 29. 

Occurrence. Dahab, 3.1.49 and 2.H.49 and 14.ii.49, shore; Sanafir, 5.ii-49. 

Remarks. The type of Duriella nigra and Row's specimen of Heteronema erecta are 
almost identical in external form though they differ in the structure of the skeleton. 
Both specimens are, however, massive and lack the digitiform processes of the type 
of H. erecta. There is also available in the British Museum collection a preparation 
from Keller's type, and comparing this with Row's specimens suggested, in the first 
place, that the only difference between Duriella nigra and Heteronema erecta lay in 
the much greater amount of sand in the fibres of the latter. The ' Manihine ' speci- 
mens, four in all, have a sufficiently general resemblance to each other, and to the 
specimens described by Keller and Row, to be considered alongside them. In these, 
two have a skeleton approximately similar to that of Duriella nigra, one is much 
more like Heteronema erecta, and the fourth is intermediate between the two. 

With seven specimens thus available for comparison it seems certain that the 
variation in the skeleton of this species (for Duriella nigra and Heteronema erecta are 
here accepted as conspecific) is similar to that shown by me (1934, figs. 18-33) f° r 
Dysidia fragilis. In other words, that according to the amount of sand present the 
skeleton will vary from clearly defined ascending fibres cored with sand, connected 
by a secondary network free of it, to a dense network in which the spongin of all 
fibres is almost entirely obscured by a heavy intake of sand, with no perceptible 
differentiation into primary (or ascending) and secondary fibres. 



172 THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 

Supporting such a view is the fact that the amount by which the fibres are im- 
pregnated with sand varies from one part to another of the skeleton of any individual 
sponge. 

The colour of the 'Manihine' specimens ranged, in formalin, from brown to a 
deep purple-brown. 

Distribution. Red Sea. 

Carterispongia clathrata (Carter) 
(For synonymy and discussion see Burton, 1934: 574.) 

Occurrence. Sherm Sheik, n.i.49 ; Mualla, 3ii.49 ; Dahab, 13 and 14.ii.49 ; Sanafir, 
9.L49 and 4.H.49 ; Sherm-el-Moiya, 3.11.49. 

Remarks. The several fragmentary specimens have the typical cavernous appear- 
ance. The skeleton differs considerably, however, from one individual to another, 
and these differences seem to offer a gradation from the typical skeleton of this 
species to that of Euryspongia lactea. It is possible, therefore, that Euryspongia 
may ultimately prove to be synonymous with Carterispongia. 

The colour of the different specimens, in formalin, ranged from fawn or brown, 
to purple, with occasional pink patches. 

Distribution. Indian Ocean; Australia; (? West Indies). 

Hircinia ramosa Keller 

Hircinia ramosa Keller, 1889: 345, pi. xx, fig. 5. 

H. schulzei Dendy, 1905: 221, pi. xvi, fig. 3. 

H. ramosa, Row, 1911: 372; Burton, 1934: 579, pi. 1, fig. 11, text-fig. 16. 

Occurrence. Sanafir, 8.L49 and 9.U.49, littoral, growing among rocks. 

Remarks.The two specimens are typical in the structure of the skeleton but show 
less of the ramose external form. One of them is low-lying and massive, with occa- 
sional ramose portions. 

The colour of the two specimens, in formalin, was fawn and brown respectively, 
in spirit it is now olive-green and brown. 

Distribution. Red Sea; Ceylon; Australia (Barrier Reef). 

Cacospongia ridleyi, sp. n 

Cacospongia cavernosa Ridley, 1884: 590; nee C. cavernosa, Autt. 

Occurrence. Abu Zabad, 11.ii.49. 

Remarks. The name Cacospongia cavernosa has been used by many authors for 
sponges from the Indian Ocean, Mediterranean, and the West Indies. Pallas (1766: 
395) appears to have been the first to use the trivial name, but his Spongia cavernosa 
is not recognizable except as one of the Keratosa. Esper's (1794: 189) 5. cavernosa, 
based on Pallas's specimen, has been inadequately re-described by Ehlers (1870: 30) ; 
and Lamarck's specimen (1813: 371) has been shown by Topsent (1930: 13) to be 
conspecific with Ciocalypta penicillus Bowerbank. Ridley (1884: 590) recorded 
specimens under Cacospongia cavernosa from the Seychelles, and it is with these 



THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 173 

that the present specimens are to be identified. C. ridleyi agrees closely with C. 
cavernosa Schmidt (as re-described by Schulze, 1879) m external form, but the 
skeleton has larger meshes and the fibres are more heavily cored with sand-grains 
and other foreign bodies. It is, however, impossible to say, in the present state of our 
knowledge, whether the sponges from Seychelles and the Gulf of Aqaba represent a 
simple variety of the Mediterranean form. As a temporary measure at least they 
are here given full specific rank. 
Distribution. Indian Ocean. 

REFERENCES 
Burton, M. 1924. The Genus Chondrilla. Ann. Mag. nat. Hist. (9) 14: 206-209. 



1924. A revision of the Sponge Family Donatiidae. Proc. zool. Soc. Lond. : 1033-1045, 1 pi. 

1926. Sponges [in] Zoological Results of the Suez Canal Expedition. Trans, zool. Soc. 

Lond. 22: 7 x - 8 3, 7 fi g s - 

1926. Stelletta purpurea, Ridley, and its variations. Ann. Mag. nat. Hist. (9) 18: 44-49. 

1928. Report on some Deep-Sea Sponges from the Indian Museum collected by R.I. M.S. 

Investigator. Part II. Rec. Indian Mus. 30: 109-138, 2 pis., 9 text-figs. 
193 1. On a collection of marine sponges mostly from the Natal coast. Ann. Natal Mus., 

4: 337-358, 1 pi., 9 text-figs. 

1934- Sponges. Sci. Rep. Gt. Barrier Reef Exped. ig28-2g, 4: 513-621, 2 pis., 33 text-figs. 

1948. The Ecology and Natural History of Tethya aurantium Pallas. Ann. Mag. nat. 

Hist. (12) 1: 122-130. 
Carter, H. J. 1883. Contributions to our knowledge of the Spongida. Ann. Mag. nat. Hist. 

(5) 12: 308-329, pis. xi-xiv. 
1886. Descriptions of Sponges from, the Neighbourhood of Port Phillip Heads, South 

Australia (contd.). Ann. Mag. nat. Hist. (5) 17: 1 12-127. 
Dendy, A. 1905. Report on the Sponges collected by Prof. Herdman at Ceylon. Rep. Pearl 

Fish. Manaar, Suppl. 18: 57-246, 16 pis. 
Dendy, A., & Row, R. W. H. 1913. The Classification and Phylogeny of the Calcareous 

Sponges. Proc. zool. Soc. Lond.: 704-813. 
Haeckel, E. 1869. Prodromus eines Systems der Kalkschwamme. Jena. Z. Naturw. 5: 236-254. 

1872. Die Kalkschwamme : eine Monographic, 2 Bd. & Atlas. Berlin. 

Hallmann, E. F. 191 7. A Revision of the Genera with microscleres included, or provisionally 

included, in the Family Axinellidae. Proc. Linn. Soc. N.S.W. 40: 634-675, 9 pis., 4 text-figs. 
Hentschel, E. 1912. Kiesel- und Hornschwamme der Aru und Kei Inseln. Abh. senckenb. 

naturf. Ges. 34: 291-448, 9 pis. 
Keller, C. 1889. Die Spongienfauna des Rothen Meeres. Z. wiss. Zool. Leipzig, 48: 311-405, 

5 pis. 

1891. Die Spongienfauna des Rothen Meeres. Z. wiss. Zool. Leipzig, 52: 294-368, 5 pis. 

Kieschnick, O. 1898. Die Kieselschwdmme von Amboina. 66 pp. Jena. 

1900. Kieselschwamme von Amboina. Denkschr. med. -naturw. Ges. Jena, 8: 545-582, 2 pis. 

Lamarck, J. B. P. A. de M. 1813. Sur les Polypiers empates. Ann. Mus. Hist. nat. Paris, 20: 

294-312, 370-386, 432-458. 

1815. Suite des Polypiers empates. Mem. Mus. Hist. nat. Paris, 1: 69-80, 162-168, 331-340. 

Lendenfeld, R. von. 1887. Die Chalineen des australischen Gebietes. Zool. Jb. 2: 723-828, 

10 pis. 
Lindgren, N. G. 1897. Beitrag zur Kenntniss der Spongienfauna des Malaiischen Archipels 

und der Chinesischen Meere. Zool. Anz. Leipzig, 20: 480-487. 
1898. Beitrag zur Kenntniss der Spongienfauna des Malaiischen Archipels und der 

Chinesischen Meere. Zool. Jb. Jena (Abt. Syst.), 11: 283-378, 4 pis. 
Michlucho-Maclay, N. 1868. Beitrage zur Kenntniss der Spongien. Jena Z. Naturw. 4: 221- 

240, 2 pis. 



174 THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 

Nardo, G. D. 1847. Osservazioni anatomiche sopra l'animale marino detto volgarmente 

Rognone di mare. Atti. 1st. veneto, 6: 267-276. 
Ridley, S. O. 1884. Spongiida. Rep. Zool. Colls. Voy. H.M.S. 'Alert', London: 366-482, 

582-630, 6 pis. 
& Dendy, A. 1886. Preliminary Report on the Monaxonida collected by H.M.S. Challenger. 

Ann. Mag. nat. Hist. (5) 18: 325-351. 

1887. Monaxonida. Rep. Sci. Res. Voy. H.M.S. 'Challenger', Zool. 20: 1-275, 51 pis. 

Row, R. H. W. 1909. Report on the Sponges collected by Mr. Cyril Crossland in 1904-5. Part I. 

Calcarea. /. linn. Soc. Lond. Zool. 31: 182-214, 2 pis. 
191 1. Report on the Sponges collected by Mr. Cyril Crossland in 1904-5. Part II. /. linn. 

Soc. Lond. Zool. 31: 287-400, 7 pis., 26 text-figs. 
Schuffner, O. Beschreibung einiger neuer Kalkschwamme. Jena. Z. Naturw., xi, (2) 4: 403- 

433, 3 Pis. 
Thacker, A. G. On collections of the Cape Verde Islands Fauna made by Cyril Crossland. Proc. 

zool. Soc. Lond.: 757-782, 1 pi., 12 text-figs. 
Thiele, J. 1903. Kieselschwamme von Ternate. Abh. senckenb. naturf. Ges., 25: 933-968, 1 pi. 
Topsent, E. 1906. Sponges recueillies par M. Ch. Gravier dans la Mer Rouge. Bull. Mus. Hist. 

nat. Paris, 12: 557-570. 
1925. £tude des Spongiaires du Golfe de Naples. Arch. Zool. exp. gen. Paris, 63: 623-725, 

1 pi., 27 text-figs. 



IV. TURBELLARIA: POLYCLADIDA 

By STEPHEN PRUDHOE 

Though comprising only three specimens, the collection is an interesting one, since 
it includes three species which apparently have not been recorded hitherto from the 
Red Sea. 

The condition of the material is satisfactory, and it has been possible to supple- 
ment existing descriptions of the three species with some new details of their structure, 
more especially of the copulatory organs. 

Lastly, a brief historical account of the poly clad fauna of the Red Sea is given, 
together with a list of the species recorded. 

Planoceridae 
Planocera crosslandi Laidlaw, 1903 

(Fig. 1) 

A young adult specimen of this species was found in the fauna associated with 
coral at Sherm Sheik, 2 February. It measures about 28 mm. in length and about 
20 mm. in maximum width, which occurs in the middle region of the body. 




Fig. 1. Planocera crosslandi. Arrangement of eyes (dorsal view) 



zoo. 1. 8. 



Aa 



176 THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 

In the structure of the copulatory organs the present specimen agrees very well 
with the original description of P. crosslandi. The posterior region of the cirrus- 
cavity bears three very large hook-like structures, one of which is attached to the 
dorsal wall and the others to the sub ventral walls of the cavity. These structures are 
directed posteriorly and lie almost entirely in the spacious male antrum. 

Planocera crosslandi has been recorded hitherto only from British East Africa. 

Leptoplanidae 

Notoplana gardineri (Laidlaw, 1904) 

(Fig. 2) 

A single individual, provisionally assigned to this species, was found under a rock 
near the low-tide mark at Sherm Sheik, 15 February. Unfortunately the specimen is 
damaged, and, as a portion of its hinder region is lost, it is not possible to determine 
the structure of the female copulatory apparatus. 

Transverse serial sections of the copulatory organs of the type-specimen of this 
species have recently been presented to the British Museum (Natural History) by 




Fig. 2. Notoplana gardineri. Arrangement of 
eyes (dorsal view). 

Dr. F. F. Laidlaw. The series is incomplete, but, so far as it has been possible to 
make out, the male copulatory apparatus of the specimen from the Red Sea is 
indistinguishable, structurally and histologically, from that of the type-material of 
N. gardineri (Laidlaw), a species known hitherto only from Ceylon. 



THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 177 

The damaged specimen is somewhat pellucid and measures about 16 mm. in length 
and about 9 mm. in maximum width. The body is more or less oval in outline. No 
tentacles have been made out. The eyes are arranged in two elongate groups (Fig. 2). 
Those in the hinder region of each group are distinctly larger than the remainder and 
probably represent the tentacular eyes present in other species of Notoplana. 

The mouth occurs about 10 mm. from the anterior extremity of the body and 
opens into the hinder region of the pharyngeal chamber. The latter measures about 
4*5 mm. in length and contains about 10 pairs of shallow lateral pockets. 

The male pore is situated at 3-5 mm. behind the mouth. As is usual in this genus, 
the ovaries and testes lie in the dorsal and ventral parenchyme respectively. The 
vasa deferentia unite to open into the proximal end of the arcuate seminal vesicle, 
which possesses a very thick coat of longitudinal and circular muscle-fibres. This 
vesicle opens, through the ejaculatory duct, into a well-developed, somewhat pear- 
shaped prostatic organ lying above the proximal end of the seminal vesicle. The 
ejaculatory duct projects well into the prostatic organ, the highly glandular epi- 
thelium of which completely invests the duct. In this epithelium there are seven 
elongate pockets, which, together with the ejaculatory duct, open into a small 
chamber situated in the posterior region of the prostatic organ. From the prostatic 
chamber a long ductus communis or prostatic canal passes through an extremely 
thick sheath of muscle-fibres and enters a very small penis-papilla lying in the 
shallow male antrum. The thick sheath appears to be a continuation of the muscula- 
ture of the prostatic organ and merges with that of the penis-papilla. There are 
numerous nuclei present among the muscle-fibres of the sheath, and they seem to 
congregate more particularly around the prostatic canal. 

N. gardineri appears to bear a very close resemblance to Notoplana otophora 
(Schmarda, 1859) which was also originally recorded from Ceylon. According to 
Stummer-Traunfels (1933), the 'ductus communis' or prostatic canal of the type- 
specimen of N. otophora is invested with a deep layer of parenchymatous tissue 
enclosed in a thick muscular sheath. On the other hand, in N. gardineri the prostatic 
canal is, as stated above, invested solely with an extremely thick musculature of 
longitudinal and circular fibres. Nuclei are abundant in this musculature, being 
particularly dense immediately around the prostatic canal. This difference between 
the two species might be accounted for by the fact that the type-specimen of N. 
otophora had, when examined by Stummer-Traunfels, apparently been stored in 
preserving fluid for about seventy years. During this time the tissues of the speci- 
men had, no doubt, undergone some maceration and possibly the histology of the 
structure through which the prostatic canal passes might originally have been 
similar to that occurring in N. gardineri. In other respects, except possibly in the 
number of eyes, the two species appear to be identical. 

Notoplana cotylifera Meixner, 1907 

A single specimen was found in sponges associated with coral at Graa, 30 January. 
It agrees very well with the description of N. cotylifera Meixner, and, as in the original 
material, a well-developed sucker occurs between the genital pores. 

The most striking feature of the female copulatory apparatus in this species is 



178 THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 

the pocket-like structure, which Meixner regards provisionally as a rudimentary 
accessory vesicle, opening into the vagina interna, near the ' shell '-chamber. A 
somewhat similar structure occurs in the present specimen, but in this instance it 
appears also to open on the dorsal surface of the body, anteriorly to the female 
genital pore. Unfortunately the condition of the tissues in this region of the body is 
not very satisfactory, and the presence of a dorsal opening requires confirmation. 
If a study of new material were to show that the dorsal opening normally occurred 
in this species, the accessory structure of the female apparatus would appear com- 
parable with the ductus vaginalis present in some other species of Polyclads. 

Notoplana cotylifera has been recorded previously from the Gulf of Tadjoura, 
French Somaliland, which is, of course, situated near the southern entrance to the 
Red Sea. Thus the occurrence of this species in the Gulf of Aqaba is not unexpected. 

The history of the Polyclad fauna of the Red Sea apparently begins in the year 
1826, when the name Planaria mulleri was given by Audouin to a planarian figured, 
but not described, by Savigny in the same year. Two years later (1828) Leuckart 
described five new forms from Tor in the Gulf of Suez. This work was shortly followed 
by that of Ehrenberg (1831), in which a further four new species were described from 
Tor and the Isle of Ras el Gusr. The descriptions of all these ten species are very 
incomplete, and it does not appear possible to recognize any of the species with 
certainty. 

After 1 83 1 no further species of Polyclads seem to have been recorded from this 
region until Boutan (1892) mentioned the occurrence of Pseudoceros violaceus 
(Schmarda) at Port Tewfik. Another thirty years elapsed before Meyer (1922) 
described three new species from Kosseir. Since the appearance of Meyer's work, 
Palombi (1928) has recorded, among other species, Idioplana australiensis Wood- 
worth 1 from the Port of Suez, and Melouk (1940, 1941) has described two new forms 
from the Biological Station at Ghardaqa. 

The results of the sporadic work done since 1826 indicate that our knowledge of 
the occurrence and distribution of Polyclads in the Red Sea is, in all probability, 
very incomplete. It may therefore be deemed useful to tabulate the species, including 
those in the present collection, that have so far been recorded from the Red Sea. 
The taxonomy of some of the species is very uncertain, and these are marked with an 
asterisk in the following table : 

Species Locality 

Cestoplana polypora Meyer, 1922. . . . Kosseir 



' Craspedomata sp. ? ' Palombi, 1928 
Cryptophallus aegyptiacus Melouk, 1940 
*Eurylepta flavomarginata Ehrenberg, 1831 
*Eurylepta praetexta Ehrenberg, 1831 

Idioplana australiensis Woodworth, 1898 
*Leptoplana hyalina Ehrenberg, 1831 

[This species, the type of the genus Leptoplana, has been regarded by most 
early writers as a synonym of Leptoplana tremellaris (Miiller, 1774).] 



Gulf of Suez 

El Ataka & Ghardaqa 

Ras el Gusr 

Tor 

Port of Suez 

Tor 



1 Judging from Palombi's description, the material determined by him as Idioplana australiensis is 
probably not identical with that described by Woodworth. In fact, Palombi's material appears to be 
more closely related to the genus I dioplanoides Barbour, 1912, than to Idioplana Woodworth, 1898. 



THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 



179 



Species 
Leptoplana nadiae Melouk, 1941 . 
Notoplana cotylifera Meixner, 1907 
Notoplana gardineri Laidlaw, 1903 
Paraplanocera marginata Meyer, 1922 
*Planaria bilobata Leuckart, 1828. 
*Planaria bituberculata Leuckart, 1828 
*Planaria gigas Leuckart, 1828 
*Planaria limbata Leuckart, 1828 . 
*Planaria miilleri Audouin, 1826 . 

[P. bituberculata and P. miilleri have been generally regarded as synonyms 
of Stylochus suesensis Ehrbg. If this be accepted, P. miilleri has priority 
over 5. suesensis and therefore becomes the type-species of Stylochus 
Ehrbg.] 
*Planaria zebra Leuckart, 1828 .... 
Planocera crosslandi Laidlaw, 1903 
Pseudoceros violaceus (Schmarda, 1859) 
Stylochus coseirensis Bock, 1927 [nom. nov. pro 
Stylochus reticulatus of Meyer, 1922] . 
* Stylochus suesensis Ehrenberg, 1831 



Locality 
Ghardaqa 
Graa 

Sherm Sheik 
Kosseir 
Tor 
Tor 
Tor 
Tor 



Tor 

Sherm Sheik 

Port Tewfik 

Kosseir 

Tor & Port of Suez 



REFERENCES 

Audouin, V. 1826. Explication sommaire des planches. Annelides. Descr. £gypte, etc. Hist. nat. 

1(4): 76. 
Boutan, L. 1892. Voyage dans la Mer Rouge. Rev. Biol. Nord France, 4: 173-183. 
Ehrenberg, C. G. 1831. Symbolae physicae, etc. [Invertebrata.] Berolini. 
Laidlaw, F. F. 1903. On the marine Fauna of Zanzibar and British East Africa, from collections 

made by Cyril Crossland in the years 1901 and 1902. — Turbellaria Polycladida. Part I. 

The Acotylea. Proc. zool. Soc. London, 2: 99-113, pi. ix. 
1904. Report on the Poly clad Turbellaria collected by Professor Herdman, at Ceylon, in 

1902. Rep. Pearl Fish. Manaar, Pt. II: 127-136, pi. 
Leuckart, F. S. 1828. In Leuckart & Ruppell: Neue wirbellose Thiere des Rothen Meeres. 

Atlas zu der Reise im nordlichen Afrika von E. Ruppel. Abth. I, Zoologie: 11 & 15, pi. iii. 

Frankfurt a. M. 
Meixner, A. 1907. Polycladen von der Somalikuste, nebst einer Revision der Stylochinen. 

Zeitschr. wiss. Zool. 88: 385-498, pis. xxv-xxix. 
Melouk, M. A. 1940. A new Polyclad from the Red Sea, Cryptophallus aegypticus nov. spec. 

Bull. Fac. Sci. Egypt. Univ., 22: 125-140, pis. i-ii. 
— — 1941. Leptoplana nadiae, a new Acotylean Polyclad from Ghardaqa (Red Sea). Bull. Fac. 

Sci. Egypt. Univ., 23: 41-49, pi. i. 
Meyer, F. 1922. Polycladen von Koseir (Rotes Meer). Arch. Naturgesch., Abt. A, 87: 138-158, 

pis. i-iii. 
Palombi, A. 1928. Report on the Turbellaria. [Zool. Results of the Cambridge Expedition to 

the Suez Canal, 1924. xxxiv.] Trans, zool. Soc. Lond., 22: 579-631, pi. i. 
Stummer-Traunfels, R. von. 1933. Polycladida (contd.). Bronns Klassen, 4 Abt. ic, (179): 

3486-3596, pi. i. 



V. GEPHYREA 

By A. C. STEPHEN 

ROYAL SCOTTISH MUSEUM 

Through the courtesy of the British Museum (Natural History), I have had the 
privilege of examining this collection. It is a small one containing seven individuals, 
referable to two genera of Sipunculids and one Echiurid. With one exception they 
have been recorded previously from the Red Sea, the exception being Siphonosoma 
koreae Sato, whose status is discussed. 

Echiuridae 

Ochetostoma erythrogrammon (Leuckart & Ruppell) 

Sherm Sheik, 15.ii.49. Under rock at low tide. One specimen, body 30 mm., 

proboscis 22 mm. 
This species has already been recorded from a number of localities in the Red Sea. 

Sipunculidae 
Siphonosoma koreae Sato 

Sherm-el-Moiya, 3.U.49. Associated with coral. One specimen, not fully extended, 
115 mm. in length. 

A single specimen, which agrees closely with Sato's description (Sato, 1939: 379), 
was secured. The body is long and thin, pink in colour, and capped at both ends by 
areas of yellow colour, the posterior area being much less extensive than the anterior 
area. The body is translucent, the muscle-bands showing through clearly. 

The posterior end of the body is somewhat cone-like, and the yellow cap extends 
for a distance of 5 mm. The introvert is not fully extended, but the yellow area 
occupies some 20 mm. of the body. 

In the specimen described by Sato the colour of the body is given as greyish 
white. 

The skin has numerous papillae, prominent and closely packed on the posterior 
end and at the base of the introvert, small and scattered on the rest of the body. 

Sato described the papillae on the posterior end in his specimen as being less 
prominent than those on the introvert basis. In this specimen, however, they are 
of similar size. On the introvert basis the area of prominent papillae extends for 
about 4 mm. 

On the introvert the papillae are small and arranged on circular ridges. 

The longitudinal muscle is divided into 19 bands, as in Sato's specimen. 

This species was described by Sato from a single specimen taken at Gunzan in 
Korea on 2 September 1937. In his key and text it is described as being very similar 
to S. cumanense (Keferstein), separable mainly by colour differences, especially the 
yellow caps, and by the character of the papillae on the basis of the introvert. The 



182 THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 

specimen from Aqaba differs from the Korean one in the colour of the body and the 
greater prominence of the posterior papillae. In view of the somewhat protean nature 
of 5. cumanense, with its three well-marked and widely distributed varieties, of which 
two are common to both the Red Sea and Korean waters, as well as the differences 
between the two known specimens, it is possible that more material may show that it 
is not a distinct species but only another variety of 5. cumanense. 

Physcosoma pacificum (Keferstein) 

Abu Zabad. 11.ii.49. On reef at low tide. Two specimens. One partially ex- 
tended, 12 cm. in length. The other similar in size but much contracted. Greyish 
brown in colour, with scattered darker patches. 

Tiran. io.i.49. Associated with coral. One large specimen; not fully extended, 
about 13 cm. in length. Uniformly greyish brown in colour with a number of 
darker bands anteriorly. 

Dahab. 3.H.49. Shore. Two specimens of similar size to the above, but too 
contracted for measurement. Greyish brown in colour, with scattered darker 
patches. 

This species is widely distributed in the Indo-Pacific area and has already been 
recorded from the Red Sea. 



REFERENCE 

Sato, H. 1939. Studies on the Echiuroidea, Sipunculoidea, and Priapuloidea of Japan. Sci. 
Rep. Tdhoku Univ. (4) 14: 339-460, 5 pis., 60 figs. 



VI. MOLLUSCA 

By w. j. rees and a. stuckey 

The mollusca are represented by 2 Loricates, 27 Gastropods, 13 Lamellibranchs, 
4 Cephalopods, and a few Nudibranchs not reported on here. As this particular 
area has been thoroughly worked for mollusca by numerous workers, notably 
McAndrew and Issel, it is not surprising that no new forms were found. The Gastro- 
poda and Lamellibranchia call for no special description and have been listed with 
notes on distribution. Although the Cephalopoda are all well-known species, they 
are so well preserved that we have noted features of interest, standard measure- 
ments, and included photographs. 

Callistochiton heterodon var. savignyi appears to be rare and is only known from 
the northern part of the Red Sea ; it was not hitherto represented in the collections 
of the British Museum. Other species which appear to be confined to the Red Sea 
are Clanculus pharaonis, Trochus erythraeus, and Lithophaga hanleyana. All the 
remaining species are found either in the western part of the Indian Ocean or have 
a wide distribution in the Indo-Pacific. In the Cypraeidae Schilder (1938) has drawn 
attention to races of well-known species which are becoming differentiated in various 
areas, including the Red Sea. 

The classification of Indian Ocean Lamellibranchs, and indeed all Lamellibranchs, 
is in a very unsatisfactory state, and in the specific names we have adopted we have 
followed Thiele (1929-1934), Tomlin (1927), Smith (1897), and various papers by 
E. Lamy. Recent work on molluscs has shown that species seemingly identical, or 
appearing to have only minor points of difference, have distinct larvae and life 
histories, revealing that they are distinct. Elaborate lists of synonyms may therefore 
prove erroneous, and usually we have confined ourselves to referring the specimens 
to species with which they appear to be identical. 

Among the Cephalopods Octopus macropus is common in the Red Sea and in the 
Mediterranean. The remainder, 0. horridus Orbigny, 0. cyanea Gray, and Sepio- 
teuthis lessoniana Lesson, are at the western limit of their range, which extends to 
the Andaman Islands in 0. horridus and throughout the tropical Indo-Pacific in the 
other species. 

The Cephalopods of the Red Sea have been reviewed by Adam (1942) and a study 
of his list reveals that they are all either littoral and shallow water forms or have 
planktonic larvae which live close to the surface during their early life. 1 As examples 
of the former we have species of Octopus, Sepia, Sepioteuthis, and Doryteuthis, and 
of the latter (oceanic species) we have Symplectoteuthis , Tremoctopus, and Argonauta. 
Cephalopods characteristic of deep water, and even the Cranchiidae (pelagic species 
which spend much of their early life in the upper 500 metres), are absent. 

It has been pointed out by Thompson (1939) that there is a shallow sill near 

1 We have excluded Spirula spirula (L.), the shells of which are recorded from the Red Sea. It is 
probable that these have their origin outside the area. 

ZOO. I. 8. Bb 



184 THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 

Hanish Islands separating the Red Sea 'proper' from the Gulf of Aden. At about 
latitude 13 41/ N. the depth of the sill is only 100 metres, and this may act as a 
geographical barrier to deep-water species. This is probably one reason why bathy- 
pelagic forms are absent in the Red Sea, but it does not explain the absence of 
Cranchiidae, which as larvae often occur right up to the surface. The normal inter- 
change of water over the sill (see Thompson) should carry the larvae into the Red 
Sea and another explanation is required. There are some grounds for believing that 
these forms are the young of little known bathypelagic species and they may not be 
tolerant to high salinities of 38% to 40% such as are typical of the Red Sea. 

The following species are known only from the Red Sea; those marked by an 
asterisk are insufficiently known and may prove to belong to other species: 

Sepia savignyi Blainville, 1827 Doryteuthis arabica Ehrenberg, 1831 

* Sepia gibba Ehrenberg, 1831 *Abralia steindachneri Weindl, 1912 

* Sepia elongata Ferussac & d'Orbigny, Octopus robsoni Adam, 1941 

1835-1848 Sepia dollfusi Adam, 1941 

*Sepia trygonina Rochebrune, 1884 

Four of the above are imperfectly known, and of the seventeen species recorded 
from the Red Sea, only four sound species can be regarded as endemic. It is possible 
that even this number may be further reduced when the cephalopod fauna of the 
western Indian Ocean becomes better known. 



Class LORICATA 

Family Crytoplacidae 

Callistochiton heterodon var. savignyi Pilsbry 

Locality: 30^.49, Mualla, 1 specimen. 

This small Callistochiton was taken with two specimens of Acanthopleura haddoni. 
It has a total length of 13 mm. and a breadth of 7-5 mm. 

This variety was named by Pilsbry (1892) from a figure given by Savigny (Egypte, 
pi. 3, fig. 8). Our specimen has the following characters. Shell oval, distinctly ridged 
along the median line; the sides of the valves are only slightly curved. Valves 
greyish white with occasional irregular darker markings. Girdle buff-coloured with 
faint slate-grey vertical bands. Head valve, with 10 slightly denticulate ribs, 2 of 
these bifurcate anteriorly. Tail valve, distinctly narrower than head valve, with 11 
radiating rays. Other valves with distinct but not backward projecting beaks. 
Lateral areas raised with 2 denticulate ribs. Central areas with 7-8 narrow deeply 
etched riblets on each side with a central smooth tract between them. 

This variety has affinities with C. adenensis Smith, but differs from it mainly in 
having only 10-11 radiating ribs on the anterior valve instead of 22 as in Smith's 
species. C. heterodon var. savignyi is only known from this northern part of the 
Red Sea. 



THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 185 

Family Chitonidae 

Acanthopleura haddoni Winckworth 

Acanthopleura sp. Haddon, 1886: 24. 

Chiton (Acanthochites) spiniger Issel, 1819: 235 [non Sowerby]. 
Acanthopleura spinigera, Sykes, 1907: 34; Tomlin, 1927: 292 [non Sowerby]. 
Acanthopleura haddoni Winckworth, 1927: 206. 

Localities: 29.xii.48, Aqaba, just below low tide mark, 4 adults. 8.1.49, Sanafir, 
2 adults. 30.1.49, Mualla, attached or under stones at low tide, 2 adults. 9.H.49, 
Sanafir, among rocks along the shore, 1 adult. 

This large and decorative chiton is known from the Suez area of the Red Sea under 
the name spinigera. The earliest record is that of Savigny (Egypte, pi. 3, fig. 4). 
Winckworth (1927) distinguishes the species from A. spinigera Sowerby, an Austra- 
lian and Indonesian species, and described specimens from Aden under the name 
A. haddoni. According to Winckworth the living animal reaches a length of 3 in. 
and our largest specimen is of this size, although it cannot be measured accurately 
because of contraction of the foot causing the animal to be bent in the form of a 
crescent. In all our examples the girdle is irregularly marked with black and olive 
bands. In the living animal the foot is of a salmon-pink colour. 

It is impossible at present to give an accurate picture of the distribution of this 
mollusc. We know it occurs in the Red Sea (at Aden, Suez, and the localities given 
above), but its occurrence outside this area becomes confused with that of A. spini- 
gera (Sowerby). Cyril Crossland (quoted by Sykes) notes that it is 'the common 
high tide chiton, everywhere in E. Africa, on the cliffs of coral-rag at Djibouti, 
Mombasa, Zanzibar, Wasin etc. ; also on stone on the edge of reefs of the East Coast 
of Zanzibar*. 

Class CEPHALOPODA 
Family Loliginidae 
Sepioteuthis lessoniana Lesson (Plate 28, figs. 1 and 2 ; Plate 29, figs. 5 and 6) 

31.xii.48, Station Ai, dip net, surface, Faraun Island, i?(3). 28.L49, Aqaba, 
i$(i). 4.H.49, Sanafir, cast net, surface, 1^(2). 

The two specimens are remarkably well preserved, and the ground colour in 
formalin is flesh-coloured. The reddish-purple chromatophores are fairly evenly 
distributed over the ventral surface of the head, arms, funnel, and mantle, with 
denser patches around the edge of the eye. There are no chromatophores on the 
ventral surface of the fins ; the muscle-fibres of the fins are prominent. 

On the dorsal surface, the chromatophores are more densely crowded, especially 
just above the eye, and on the dorsal mantle. 

In the male (less prominent in the female) there are irregular ivory-coloured 
patches which are covered by one or more large chromatophores. When the skin of 
the mantle is folded back to expose the pen, these white patches are seen to lie 
over patches of bright emerald-green, which presumably cause the animal to be 
iridescent. The secondary sexual character of the male (transverse whitish streaks 
across the dorsal mantle), which has been noted by Adam (1938), is distinct in the 



i86 



THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 



large male from Sanafir (No. 2). Colour notes on the living animal state that 
the dorsal mantle was of a reddish-brown colour with iridescent green spots on the 
mantle itself, but not on the fins. The animals appear to fall within the limits given 
by Adam (1939) in his review of Sepioteuthis lessoniana. Adam, and indeed most 
other workers, have given figures of medium-sized individuals of about 150 mm. in 
dorsal mantle length. In these the maximum fin width is found in the posterior 
third of the body, but in our specimens, which are much larger, the maximum fin 
width occurs about midway between the apex and the mantle margin. This is to be 
expected in the larger animals because growth proceeds at a much faster rate in the 
posterior third of the body. As far as can be judged from Adam's illustrations of S 
hemprichii Ehrenberg 1831 our specimens agree in form of the body and in the fins. 
There appears to be no doubt that Ehrenberg's specimens were really large indi- 
viduals of 5. lessoniana like these from the Gulf of Aqaba. The hectocotylized arm 
(left ventral arm) is particularly well developed and of the usual pattern in Sepio- 
teuthis. There are 34 pairs of suckers which gradually diminish in size distally, with a 
proportionate increase in size of their peduncles. The distal portion of the arm is 
occupied by 25 pairs of triangular flattened papillae. As noted by Adam (1939) the 
papillae on the dorsal side are more strongly developed than those on the ventral side. 
In the female specimen spermatophores have been deposited on the ventral side 
of the buccal membrane. 

Table I 

Sepioteuthis lessoniana Lesson 

(Measurements in mm.) 





(1)? 


(2)3 


(3)? 


Dorsal mantle length ...... 


280 


270 


136 


Ventral mantle length 












260 


235 


127 


Greatest mantle length 












65 


80 


35 


Greatest mantle thickness 












60 


63 


34 


Length of head 












39 


61 


31 


Width of head 












64 


71 


36 


Thickness of head . 












45 


46 


23 


Length of fin . 












258 


246 


124 


Distance between fin base and mantle margin 






5 


6 


6 


Arms 








1st right ........ 


72 


87 


30 


1 st left 


















69 


80 


28 


2nd right 


















102 


in 


47 


2nd left 


















39 (broken) 


105 


5i 


3rd right 


















128 


131 


58 


3rd left 


















64 (broken) 


122 


60 


4th right 


















122 


136 


62 


4th left 


















66 (broken) 


134 


62 


Length, right tentacle 














197 


214 


95 


Length, left tentacle 














71 (broken) 


244 


92 


Right tentacular club 














no 


101 


40 


Left tentacular club 














(missing) 


116 


42 


Diameter largest arm sucker 












5 


5 


2'5 


Diameter largest tentacular sucker 










7 


8 


4 



THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 



187 



Sepioteuthis sp. (Plate 29, figs. 3 and 4) 
i.ii.49, Sherm Sheik, surface, 1 juvenile, n.i.49, Sherm Sheik, 1 newly hatched. 

The young post-larval squid compares very favourably with one illustrated by 
Wiilker (1913, pi. 22, fig. 2g). In our specimen the chromatophores are more numer- 
ous than in Wulker's slightly younger specimen. Full measurements of this specimen 
are given in Table II. We are not able to assign this to any particular species of 
Sepioteuthis, but if we may judge by the extent to which the fins are developed 
there is every likelihood that it is a young individual of Sepioteuthis lessoniana 
Lesson. 

The newly hatched larva has a dorsal mantle length of only 4-5 mm. and is a little 
damaged. It compares very favourably with a stage illustrated by Wiilker in his 
figure 2g. 

Table II 

Sepioteuthis sp. 



(Measurements in mm. 



Dorsal mantle length .... 
Ventral mantle length .... 
Greatest mantle breadth 
Greatest mantle thickness 
Length of head 

Width of head 

Thickness of head ..... 

Length of fin 

Distance between fin base and mantle margin 



19 
17 

7 

6-5 

7 

7 

6 
12 

5 



irms 


Right 


Left 


1st . 


• 3-5 


3'5 


2nd . 


• 7 


7 


3rd . 


. 10 


10 


4th . 


. 8 


8 



Length, right tentacle 

Length, left tentacle 

Right tentacular club 

Left tentacular club 

Diameter of largest arm sucker 

Diameter of largest tentacular sucker 



16 

16 
7-5 
7-5 
o-35 
0-4 



Octopus horridus d'Orbigny (Plate 29, fig. 7) 

Octopus horridus d'Orbigny, 1826: 144. 

Octopus argus Krauss, 1848: 132. 

Polypus aculeatus Hoyle, 1904: 194 [non d'Orbigny 1840]. 

Octopus {Octopus) horridus, Robson, 1929: 91. 

io.i.49, Tiran, found in coral, 1 <J. 

This littoral octopus is well known from the Suez area of the Red Sea. It has been 
previously taken in the crevices of coral by Hoyle (1907). 

Our specimen agrees in most particulars with earlier descriptions, but a few features 
are worthy of comment. The dorsal surface of the mantle, head, and arms is orna- 
mental with pale olive-green patches; most of these have a distinct cirrus in the 



i88 



THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 



centre. The spaces in between the paler patches are rilled by closely grouped chroma- 
tophores, which appear black or very dark red in formalin. There is no ocellus. 
The ventral surface of the mantle is of a pale cream colour. Colour notes on the 
living animal state that when found the Octopus was yellowish with a green network 
on the arms. The ground colour changed to brown when the animal was placed on a 
dark background. 

The ground colour of the inner surface of the tentacles is also pale cream with light 
brown chromatophores evenly distributed over it. 

The body is ovoid, the eyes prominent, and the arms long in proportion to the 
length of the body (the arms are too tightly coiled for accurate measurements, but 
the formula is of the order 4.3.2.1). The ventral arms are more robust than the 
others, the first pair being the least well developed. As noted by Robson (1929) 
the hectocotylized arm is shorter than its fellow. The spermatophore groove on the 
ventral side is prominent, and is protected especially near its tip by a membraneous 
extension of the arm. 

The standard measurements are given in Table III. 

Distribution. This species has been recorded by a number of workers, from the 
Red Sea, and especially from the Suez Canal zone (see Robson, 1929). Beyond the 
Red Sea it has been recorded from Ceylon, and other parts of the central Indian Ocean 
by Hoyle (1904, 1905, 1907a and b). Other records from the same area are given by 
Robson (1929: 91). There are no records of this species east of the Andaman Islands. 



Table III 

Octopus horridus d'Orbigny 

(Measurements in mm.) 



Sex 

Total length (including 3rd arm) 

Dorsal mantle length . 

Width of body . 

Width of head . 

Arm formula 

Web formula 

Diameter of largest sucker 

Length of ligula . 



D> C 



= E 



• 65 + 

• 15 
. 12 



4.3.2.1 
> B> A 

. 2-25 
• 2-55 



Indices 

Mantle width index . 
Head width index . 
Sucker index (normal) 



. 80 
. 13 5 
. 15 



Octopus macropus Risso 

11.ii.49, Abu Zabad, on reef at low tide, i $. 31.xii.48, Station Ai, Faraun Island, 
surface, imm. $. 

This well-known octopus needs no further description, but standard measure- 
ments are provided for comparison with those which already exist for the Caribbean 
population of this species (Table IV). The measurements indicate that the Red 
Sea specimens fall within the limits already known for the species. 



THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 189 

Distribution. The species occurs in the Caribbean, the NE. Atlantic, the Mediter- 
ranean, the Red Sea, and the Indo-Pacific to Japan and Australia. Its eastern limit 
appears to be the Marshall Islands. It has been recorded from the Red Sea by Wiilker 
(1920) and Weindl (1912), to mention only two records. 



Table IV 
Octopus macropus Risso 





(Measurements in mm.) 






Sex 






? 




juvenile <£ 


Total length (including 3rd arm) . 




246 






40 


Dorsal mantle length . 






58 






16 


Eye to dorsal web 






47 






6 


Width of body . 




36 


36 






10 


Width of head 




— 


29 






7 


Arms 




Right 




Left 


Right 


Lej 


1st . 




246 




245 


34 


34 


2nd. 




228 




227 


28 


26 


3rd . 




208 




177 


23 


22 


4 th . 




186 




190 


20 


20 


Diameter of largest sucker 




— 


6 






o-75 — 


No. of gill filaments . 






— 






11 


Web formula . 




. B > A 


> C> 


D> E 


B > C 


= A = D - 


Indices 














Mantle width index . 




. . 


. 62 






625 


Head width index 




. 


. 50 






44 


Sucker index (normal) 




. 


. 105 




4-7 


Arm length index 




. 


. 78-5 




68 



Table V 

Octopus cyanea Gray 

/ 11 

Sex $ ? 

Total length (including longest arm) .... 420 343 

Dorsal mantle length ....... 52 55 

Eye to dorsal web ....... — 44 

Width of body 46 3« 

Width of head 40 35 

Arms Right Left 

1st — 265 205* 

2nd ......... — | 280 190 

3rd ......... — 200* 260J 

4th — 190J 210J 

Diameter of ocellus ....... 8 5 

Diameter of largest sucker ..... 6 5 

No. of gill filaments. ...... 7-8 9 

Web formula — D = C>B>A=E 

Arm formula ........ — 2.1.3.4 or I2 -3-4 

Web depth ........ — 47 

* Arm incomplete, tip portion missing. f Arms too tightly coiled for accurate measurements. 
X Regenerating. 



i 9 o THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 

Indices 

Mantle with index 88-5 69 

Head width index 77 63-5 

Sucker index (normal) . . " . . . . 11-5 9-1 

Arm length index 82 81-5 

Web depth index — 16*8 



Octopus cyanea Gray (Plate 30) 

Octopus cyanea Gray, 1849: 15. 
Octopus marmoratus Hoyle, 1886: 227. 
Octopus horsti Joubin, 1898: 23. 
Polypus fontanianus Robson, 1920: 437. 
Polypus horsti, Wiilker, 1920: 51. 

6.L49, Sanafir, along shore, 1 $. 12.L49, Sherm Sheik, in shallow water along 
shore, 1 ?. 

We have referred these two specimens to Octopus cyanea Gray, but as they present 
a different appearance to what is usually associated with 0. cyanea, the various 
features worthy of note are discussed below. Typical specimens, of which we have 
seen a number in the collections of the British Museum, are, as Robson says, 'mainly 
of a warm ochreous red suffused and maculated with purple, which may be very deep 
so as to render the animal homogeneously blackish or deep livid (in preservative) \ 
Our specimens, however, are of a buff or pale brownish colour, with an olive-green 
sheen, which is especially marked on the dorsal surface of the web and the base of 
the tentacles. The top of the head, between the eyes, is a deeper brown colour. The 
specimens are paler ventrally and the ventral side of the arms have the character- 
istic zebra-like marking which Robson regards as one of the most striking and 
constantly associated features of cyanea as a species. Colour notes made from the 
living animal state that the colour of the specimen taken on I2.i.49 was brown and 
that the zebra-like markings on the arms were of a light blue colour. 

The dark purple ocellus is well marked and surrounded by an ill-defined pale ring, 
as mentioned by Robson for his British Museum specimens (Nos. 4 and 8). 

Specimen No. I is rather contracted ; the skin of the mantle is reticulated and has 
a number of scattered irregularly arranged cirri, which are more numerous between 
the eyes and on the fore part of the head. Specimen No. II is less contracted, and 
has four cirri arranged in a diamond pattern on the dorsal mantle and four to five 
prominent cirri on the fore part of the head. The ventro-lateral and anterior portion 
of the mantle carries a number of scattered cirri. There is also a curious fold of skin, 
on either side of the neck region postero-ventral to the eye, which effectively separates 
the ventral funnel region from the lateral face of the head. 

The dorso-lateral surface of the arms in both specimens have a double row of 
slightly raised, buff-coloured, simple papillae which have not been mentioned by any 
other writer. A re-examination of Gray's type of 0. cyanea and other specimens 
reveals the presence of these papillae, but they are more difficult to see than in our 
specimens, because they are obscured by the dark, ground colour normal in this 
species. 



THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 191 

The number of gill filaments in the demi-branchs, normally a good diagnostic 
feature in octopods, appears to be rather variable in the species (7-9 in our speci- 
mens). Robson gives 9-10 for Gray's type of 0. cyanea, and we have found that even 
in the same specimen one gill may have 7 filaments and the other 9 filaments per 
demi-branch (1 $ from the Cocos-Keeling Islands). 

Standard measurements are given in Table V, but it has not been possible to give 
measurements of the arms in specimen I because they are too tightly coiled. 

The only other ocellate species recorded from this area is Octopus robsoni Adam, 
1941, of which a complete description has not yet been published. Adam states that 
this octopod 'se caracterise a premiere vue par la presence d'une paire d'ocelles 
pourvue d'un anneau irise blanchatre, bleuatre ou mauve '. We have mentioned this 
species because our specimens approach nearer to it in colour and the arrangement 
of the cirri than to the typical form usually found in 0. cyanea. However, the 
character of the ocellus, without an iridescent ring, the zebra-like markings on the 
ventral surface of the arms, and the various indices which fall within the limits of 
O. cyanea, leaves us in no doubt as to the identity of our species. 

Distribution. Octopus cyanea is a littoral species well known as a reef-inhabiting 
octopod, with a distribution ranging through the Indo-Pacific in tropical and sub- 
tropical waters from Hawaii to the Red Sea. 

Previous records from the Red Sea are given by Robson (1929) and Wiilker (1920). 

Class GASTROPODA 

Family Haliotidae 

Haliotis varia L. 

31.xii.48, station Ai, shore of Faraun Island, 3 specimens. 20.1.49, Dahab, on mud 
flats at low tide, i specimen. 5.H.49, Sanafir, found on coral, 1 specimen. 11.ii.49, 
Abu Zabad, on reef at low tide, 4 specimens. 15.ii.49, Sherm Sheik, under rocks at 
low tide, 1 specimen and 1 juvenile. Dahab, found on coral, 1 specimen. 

Issel (1869) collected two specimens of H. varia from the Gulf of Suez. From the 
numbers obtained in our collection it appears to be fairly common in the Gulf of 
Aqaba. According to Pilsbry (1890) it has a wide distribution in the Indo-Pacific, 
being found in the following places: Australia and Philippines to China; Mozam- 
bique, Red Sea, Island of Bourbon, Mauritius, Ceylon, Nicobar Islands, Malay 
Archipelago. 

Family Fissurellidae 

Diodora ruppellii (Sowerby) 

Fissurella ruppellii Sowerby, 1838: 128. 
Fissurella costaria Vaillant, 1865: 109. 
Fissurella vaillanti Fischer, 1865: 244. 
Glyphis ruppellii, Pilsbry, 1890: 217, pi. 39, fig. 8. 
Diodora ruppellii, Tomlin, 1927: 289. 

15.ii.49, Sherm Sheik, under rock at low tide, 1 specimen. 

Distribution. This molluscs seems to be common almost throughout the Suez 

zoo. 1. 8. cc 



192 THE 'MANIHINE* EXPEDITION TO THE GULF OF AQABA 

Canal according to Tillier & Bavay. It has frequently been reported at Suez (see 
Tomlin, 1927, for previous records). D. ruppellii is found in the Western Indian Ocean, 
in the Red Sea, at Aden, Mauritius, and on the East African coast. 

Family Patellidae 

Cellana rota (Gmelin) 

Patella rota, Issel, 1869: 233. 
Patella rota, McAndrew, 1870: 444. 
Patella variegata Reeve, 1842, pi. 136, fig. 1. 
Cellana rota, Tomlin, 1927: 299. 

12.L49, Sherm Sheik, 6 specimens. 20x49, Dahab, on mud flats at low tide, 
2 specimens. 

Both McAndrew and Issel record this species as common; the former from the 
Gulf of Suez and the latter from the Gulf of Aqaba. Tomlin (1927) found it in the 
Suez Canal zone. 

Distribution. Red Sea, east coast of Africa, Reunion, and Madagascar. 

Family Trochidae 

Clanculus pharaonis (L.) 

30J.49, Mualla, among rocks and coral at low tide, 1 specimen. 

This is one of the most characteristic molluscs of the Red Sea area ; it occurs from 
Suez to Aden, and was reported by Issel (1869) to be especially common in the Gulf 
of Aqaba. Tomlin (1927) gives previous records for the Suez area and records it 
from the Canal. 

Trochus (Infundibulops) erythraeus Brocchi 

20.1.49, Dahab, on mud flats at low tide, 1 specimen. 2.H.49, Sherm Sheik, asso- 
ciated with coral, 2 fms., 1 specimen. 

T. erythraeus has been collected from the Gulf of Aqaba by Issel (1869). Tomlin 
(1927) recorded it from the Gulf of Suez, and various other collectors, e.g. McAndrew 
(1870) and Vaillant (1865), have recorded it from the Red Sea area. 

Trochus dent at us Forskal 

30^.49, Mualla, among rocks and coral at low tide, 2 specimens. 2.U.49, Sherm 
Sheik, associated with coral, 1 young specimen. 

T. dentatus is one of the common molluscs of the Red Sea and Persian Gulf. It 
has been recorded from the Gulf of Suez by McAndrew, Issel, and Vaillant. Tomlin 
(1927) reports it from the Suez Canal zone, and Issel (1869) states that it is abundant 
in the Gulf of Aqaba. 



THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 193 

Family Turbinidae 
Turbo radiatus Gmelin 

6.H.49, Sanafir, found in coral, 1 specimen. 11.ii.49, Abu Zabad, on reef at low tide, 
2 specimens. 

T. radiatus is a common Indo-Pacific form, which is found in the Red Sea, the 
East African coast, and eastwards as far as the Philippines and New Caledonia. 
Tillier & Bavay (1905) and Tomlin (1927) record it from the Gulf of Suez and the 
Suez Canal zone. 

Family Neritidae 
Nerita forskalii Recluz 

6.i.49, Sanafir, along shore of anchorage, 3 specimens. 12.L49, Sherm Sheik, 7 
specimens. 30.1.49, Mualla, found at low tide among rocks and coral, 2 specimens. 

This extremely variable mollusc has been recorded from the Gulf of Aqaba by 
Tomlin (1927) and Issel (1869). It * s a common Indo-Pacific form, Tryon (1888) 
giving its distribution as the Red Sea, Indian Ocean, Natal, Singapore, China, the 
Philippines, and Viti Islands. 

Nerita undata var. quadricolor Gmelin 
12.L49, Sherm Sheik, 1 specimen. 

N. undata is a widely distributed species in the Indo-Pacific. In the variety 
quadricolor the aperture of the shell is white and the ribs are maculated with purplish 
black. This variety is confined to the western part of the Indian Ocean. 

Family Planaxidae 

Planaxis breviculus Deshayes 

6.L49, Sanafir, along shore of anchorage, 3 specimens. 

This species has been reported from the Gulf of Suez by McAndrew (1870), who 
records it as a common species at low water. Smith (1891) reports it from Aden and 
refers to specimens in the British Museum from the Gulf of Aqaba and Persian Gulf. 
According to Tryon (1887) P. breviculus is a variety of P. sulcatus. Both forms have 
a wide distribution in the Indo-Pacific. Until more is known about the life-history 
of these periwinkles, we prefer to retain the name P. breviculus. 

Family Cerithiidae 
Cerithium tuberculatum (L.) 
6.L49, Sanafir, shore of anchorage, 2 specimens. 

McAndrew found this species moderately common in the Gulf of Suez. It is an 
extremely variable species, and has been reported on numerous occasions from the 
Red Sea. 

Distribution. Widespread in the Indo-Pacific (Smith, 1903). 



194 THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 

Family Melanellidae 

Melanella sp. 

io.i.49, Tiran, 1 specimen. 

We do not feel justified in giving this specimen a name in view of the confusion 
which exists in the classification of the genus. 

Family Strombidae 

Pterocera lambis (L.) 

5.U.49, Sanafir, in coral, 1 specimen. 

This large shell was previously recorded from the Gulf of Aqaba by Issel (1869). 
Distribution. Widespread in the Indo-Pacific. 

Family Naticidae 
Natica matnilla L. 

N. mamilla, Lamarck, 1838: 630. 

6.1.49, Sanafir, along shore of anchorage under rocks, 1 specimen. 

N. mamilla has been previously recorded from the Gulf of Aqaba by Issel (1869). 
Tryon (1886) gives the distribution as the East Indies, the Philippines, New Cale- 
donia, and central Polynesia. 

Family Cypraeidae 

Cypraea caurica (L.) 
20.1.49, Dahab, on mud flats at low tide, 1 young specimen. 

Schilder (1938) recognizes seven races of this species, which has a widespread 
distribution in the Indo-Pacific. 

Cypraea arabica L. 

30^.49, Mualla, among rocks and coral at low tide, 1 specimen. 5.H.49, Sanafir, 
found in coral, 1 juvenile specimen, n.ii.49, Abu Zabad, on reef at low tide, 1 
specimen. 11.ii.49, Abu Zabad, on reef at low tide, 4 juvenile specimens. 

C. arabica is a well-known Indo-Pacific species, often recorded by workers on Red 
Sea fauna. Savigny (Egypte) gives a figure, and the species is recorded from the Gulf 
of Aqaba by Issel (1869). Schilder (1938) recognizes six races in the Indo-Pacific; 
our specimens conform to the E. African and Red Sea form which Schilder calls 
immanis. 

Cypraea Isabella L. 

Turia (Basilitrona) Isabella, Schilder, 1938: 176. 

3.ii.49, Sherm-el-Moiya, associated with coral, 1 specimen. 6.H.49, Sanafir, associ- 
ated with coral, 1 specimen. 



THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 195 

C. Isabella, of which four races are recognized by Schilder, has a widespread distri- 
bution in the Indo-Pacific. Our specimens belong to the typical form which is con- 
fined to the Western Indian Ocean and the Red Sea. 

Cypraea carneola L. 

Cypraea (Lyncina) carneola, Schilder, 1938: 188. 

11.ii.49, Abu Zabad, on reef at low tide, 3 specimens, n.ii.49, Abu Zabad, on reef 
at low tide, 2 juvenile specimens. 

This species is widely distributed in the Indian Ocean and also in the Pacific as far 
as Hawaii. Schilder recognizes four races of this species. The Red Sea form crassa is 
also found in the Gulf of Aden, Persian Gulf, and Karachi. 

Cypraea erosa L. 

Erosaria (Erosaria) erosa, Schilder, 1938: 137. 

30^.49, Mualla, among coral at low tide, 1 specimen. 

This species has been recorded from the Gulf of Aqaba by Issel (1869). C. erosa 
has a wide distribution in the Indian Ocean and in the Western Pacific. Our speci- 
men belongs to the typical form. Schilder (1938: 137) recognizes six races in the 
Indo-Pacific. 

Cypraea tigris L. 

Cypraea {Cypraea) tigris, Schilder, 1938: 186. 

11.ii.49, Abu Zabad, on reef at low tide, i immature specimen. 

Issel (1869) reports this species to be abundant in the Gulf of Aqaba. It has 
previously been reported from the Red Sea by many writers, including Ehrenberg 
(1831). Our specimen is not fully grown and we are unable to determine whether it 
belongs to the typical form. Cypraea tigris (sensu lata) is widely distributed in the 
Indian Ocean and in the Pacific. 

Family Cymatiidae 
Cymatium rubeculum (L.) 

Tritonium (Simpulum) rubeculum, McAndrew, 1870: 434. 
Triton (Simpulum) rubecula, Tryon, 1881: 12. 

i.ii.49, Sherm Sheik, associated with coral, 2 specimens. 

McAndrew took 2 specimens at Jubal Island in the Gulf of Suez. 
Distribution. Red Sea to the Philippines. 

Distortrix anus (L.) 
Triton anus, Reeve II, Triton, pi. xii, fig. 63. 
Abu Zabad, on reef at low tide, 1 specimen. 
This species has been previously recorded from the Gulf of Aqaba by Issel (1869). 



196 THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 

Family Muricidae 
Drupa (Drupa) ricinus (L.) 

30.1.49, Mualla, among rocks and coral at low tide, 4 specimens. 11.ii.49, Abu 
Zabad, on reef at low tide, 1 specimen. 

Distribution. Red Sea, east coast of Africa, to Natal, Philippines, and Polynesia 
(Tryon, 1880: 184). 

Drupa (Drupa) elata (Blainville) 
2.ii.49, Sherm Sheik, 2 fms., associated with coral, 3 specimens. 

This well-known inhabitant of coral reefs has a wide distribution in the Indo- 
Pacific. It is recorded from Aden by Smith (1891). 

Family Buccinidae 
Pisania ignea Gmelin 
2.ii.49, Sherm Sheik, 2 fms., 1 specimen. 5.H.49, Sanafir, found in coral, 1 specimen. 
Distribution. Red Sea, Singapore, and Philippines. 

Family Conidae 

Conus rattus Lamarck 

Conus rattus, Smith, 1891 : 399. 
Conus rattus, Dautzenberg, 1937. 

Conus rattus is a very variable species and has been recorded by many authorities 
including Smith (1891) and Dautzenberg (1937). Its distribution is very widespread 
in the Indo-Pacific. 

Conus textile L. 

11.ii.49, Abu Zabad, on reef at low tide, 1 specimen. 

This poisonous cone shell is widely distributed in the Indo-Pacific and has been 
recorded from the Gulf of Aqaba by Sturany. Dautzenberg (1937) gives a very long 
list of localities for the species. 

Family Nassidae 
Nassa pulla L. 

20.i.49, Dahab, collected on mud flats at low tide, 5 specimens. 

Issel (1869) records this shell from the Red Sea area. Tryon (1882) gives its 
distribution as the Red Sea, Java, and the Philippines. 



THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 197 

Class LAMELLIBRANCHIA 

Family Arcidae 

Area divaricata Sowerby 
Area divaricata, Tomlin, 1927: 304. 

2.ii.49, Sherm Sheik, associated with coral, 2 fms., 2 specimens. 15.ii.49, Sherm 
Sheik, under rocks at low tide, 3 specimens. 

It has previously been recorded by Tomlin from the Suez Canal and by McAndrew 
from the Gulf of Suez, under the name A. plicata. A. divaricata has a wide distribu- 
tion in the Indian and Pacific Oceans. 

Area (Barbatia) decussata Sowerby 

31.xii.48, station Ai, shore of Faraun Island, 1 specimen. 31.xii.48, station Ai, 
shore of Faraun Island, 2 specimens. 20.i.49, Dahab, mud flats at low tide, 4 speci- 
mens. 30.1.49, Mualla, among rocks and coral at low tide, 1 specimen. 9.H.49, 
Sanafir, among rocks on shore, 1 specimen. 

This species is known from the following places, according to Lamy (1917), Djibouti, 
Obock, Perim, and Aden. It is expected to have a much wider distribution, and we 
note a specimen in the British Museum collections from the Java Sea (off Batavia) . 

Family Mytilidae 
Brachidontes variabilis (Krauss) 

Mytilus variabilis Krauss, 1848: 25. 

Mytilus pharaonis Tillier and Bavay, 1905: 177. 

Mytilus exustus, Vaillant, 1865: 114. 

20.1.49, Dahab, on mud flats at low tide, 1 specimen. 

This very common species was first described from Table Bay by Krauss, who 
drew attention to its similarity to specimens from the Red Sea. The earliest record 
from the latter locality is that of Savigny (Egypte, pi. xi, fig. 5). 

Lithophaga hanleyana Reeve 
31.1.49, Mualla, associated with coral, 2 specimens. 

L. hanleyana has been previously recorded from the Gulf of Aqaba by Sturany 
(1899), who also recorded it from the Gulf of Suez and the Red Sea generally. It has 
also been recorded from the Gulf of Suez by Reeve and McAndrew. The Cambridge 
expedition to the Suez Canal (1924) also took the species in association with coral. 

Lithophaga moluccana Hanley 
14.ii.49, Dahab, associated with coral, 1 specimen. 
We have identified this species with Hanley 's species from Malacca. It appears 



i 9 8 THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 

to differ from L. hanky ana (which is already known from the Red Sea) by the more 
tapering posterior part of the shell. 
Distribution. Indian Ocean. 

Family Vulsellidae 
Vulsella vulsella (L.) 

V. lingatula, Issel, 1869: 99. 

V. mylitina, Issel, 1869: 100. 

V. trita Reeve, 1858, pi. 2, fig. 17. 

14.ii.49, Dahab, associated with coral, 1 specimen. 

Smith (191 1), who has reviewed the genus, gives the distribution of this species as 
widespread in the Indian Ocean and eastwards to Japan, N. Australia, and New 
Caledonia. From the Red Sea it has been figured by Savigny (Egypte, pi. xiv, figs. 
1 and 2) Ruppell records it as mytilina and Reeve as trita t both from the Red Sea. 

Family Pectinidae 

Chlamys luculentus (Reeve) 
Pecten luculenta Reeve, 1853, pi. 16, fig. 59. 

2.H.49, Sherm Sheik, 2 fms., associated with coral, 1 specimen. 

We have compared this specimen with the holotype of Reeve from NW. Australia 
and also with some specimens in the British Museum collection from Aden. There are 
no differences to be noted in our shell. 

The known distribution is the Red Sea and Indian Ocean. 

Family Ostreidae 
Ostrea cucullata Born 
9.H.49, Sanafir, among shore rocks, 1 specimen. 

0. cucullata is a very variable species and had been recorded from the Gulf of Suez 
by Vaillant (1865) and by Issel (1869). This oyster is edible and according to Jous- 
seaume, as quoted by Lamy (1925), is an excellent purgative. 

Distribution. Very common at many points in the Red Sea, attached to rocks, 
which are uncovered by the tide. This species is common throughout the Indian 
Ocean, and in the Pacific as far as Japanese waters (Lamy, 1925). 

Family Carditidae 
Cardita variegata (Sowerby) 

Cardium variegatum Sowerby, 1841: 107. 
Cardita subaspersa Lamarck, 1819: 25. 
Cardita radula Reeve, 1843: 191. 

n.ii.49, Abu Zabad, on reef at low tide, 4 specimens. 

C. variegata is widespread in the Indo-Pacific, Red Sea, and Australian waters. 
Lamy (1916) records it from Suez, Massaouah, Djibouti, and Perim. 



THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 199 

Family Tridacnidae 

Tridacna noae (Roding) 

Tridacnes noae Roding, 1798: 171. 
Tridacna elongata Lamarck, 1819: 106. 

31.xii.48, shore of Faraun Island, 2 specimens. 

This Tridacna has been recorded from the Red Sea, from Suez, and the Gulf of 
Aqaba by Issel (1869) under the name T. elongata Lamarck. Savigny gives the earliest 
figure from this area (Egypte, pi. x, fig. 1). It has a wide range in the Indo-Pacific, 
including Zanzibar, Mauritius, Australia, Solomon Islands, Carolines, Marshall, and 
Loo Choo Isles (McLean, 1947). 

Tridacna squamosa Lamarck 

One specimen of this common Indo-Pacific form was collected ; the label appears 
to have been lost. 

Distribution. Indian Ocean, Indonesia, Australia, the Philippines, and Japan. 

Family Veneridae 

Circe script a (L.) 
Venus scripta L. 

20.i.49, Dahab, mud flats at low tide, 1 specimen. 

Sowerby gives the distribution of this as the Red Sea and Australia. According to 
Issel (1869) it is a rare species at Suez. 



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200 THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 

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1892. Ibid. (2) 8. 

Reeve, L. A. 1841. Conchologia Systematica. 2 vol. London. 

1 843-1878. Conchologia Iconica, 20 vol. London. 

Robson, G. C. 1929. A Monograph of the Recent Cephalopoda. Pt. I. Octopodinae: 1-236, 

89 text-figs. 
Schilder, F. A., & Schilder, M. 1938. Prodrome of a monograph on living Cypraeidae. Proc. 

Malac. Soc. Lond. 23: 1 19-231, 3 text-figs. 
Smith, E. A. 1891. On a collection of marine shells from Aden. Proc. zool. Soc. Lond. 1891: 

370-436, 1 pi. 
1903. A list of species of Mollusca from South Africa forming an appendix to G. B. Sowerby's 

'Marine Shells of South Africa'. Proc. Malac. Soc. Lond. 5: 354-402, 1 pi. 

191 1. On recent species of Vulsella. Proc. Malac. Soc. Lond. 9: 306-312, 1 pi. 

Sturany, R. 1899. Catalog der bisher bekannt gewordenen Sudafrikanischen Land- und 

Siisswasser-Mollusken mit besonderer Berucksichtigung des von Dr. Penther gesammelten 

Materiales. Denkschr. A had. Wiss. Wien 67: 537-642, 3 pis. 
Sykes, E. R. 1907. Reports on the Marine Biology of the Sudanese Red Sea — V. On the Poly- 

placophora or Chitons. /. linn. Soc. Lond. 31: 31-34. 
Thompson, E. F. 1939. Chemical and physical investigations. The exchange of water between 

the Red Sea and the Gulf of Aden over the ' sill '. Sci. Rep. John Murray Exped. 1933-34. 

2: 105-119, 10 text-figs. 
Tillier, L., & Bavay, A. 1905. Les Mollusques Testac6s du Canal de Suez. Bull. Soc. zool. Fr. 

30: 170-181. 
Tomlin, J. R. le B. 1937. Catalogue of Recent Fossil Cones. Proc. Malac. Soc. Lond. 22: 

205-330. 
1927. Report on the Mollusca (Amphineura, Gastropoda, Scaphopoda, Pelecypoda). 

Trans, zool. Soc. Lond. 22: 291-320. 
Tryon, G. W. 1880. Man. of Conchology (1) 2. 
1881. Ibid. (1) 3. 



THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 201 

Tryon, G. W. 1882. Man. of Conchology (1) 4. 

1886. Ibid. (1) 8. 

1887. Ibid. (1) 9. 

1888. Ibid. (1) 12. 

Vaillant, L. 1865. Recherches sur la faune malacologique de la baie de Suez. /. Conchyliol. 

13: 97-121. 
Weindle, T. 1912. Vorlaufige Mitteilungen iiber die von S. M. Schiff 'Pola' im Roten Meer 

gefundenen Cephalopoden. Anz. Akad. Wiss. Wien, 49: 270-275. 
Winckworth, R. 1927. New species of Chitons from Aden and South India. Proc. Malac. Soc. 

Lond. 17: 206-208, 2 pis. 
Wulker, G. 1913. Cephalopoden der Aru- und Kei-Inseln. Abhandl. Senckenb. Naturf. Ges. 

34: 451-487; 1 pi., 8 text-figs., 1 sketch-map. 
1913. tiber das Auftreten rudimentarer akzessorischer Nidamentaldrusen bei mannlichen 

Cephalopoden. Zoologica, Stuttgart 26: 201-210, 1 pi. 
1920. tlber Cephalopoden des Roten Meeres. Senckenbergiana, Frankfurt, 2: 48-58. 



Legends to Plates 28-30 

PLATE 28. SEPIOTEUTHIS LESSONIANA LESSON 

Fig. 1. Ventral view of $ caught at the surface off Faraun Island, 
31.xii.48. 

Fig. 2. Dorsal view of 6* caught off Sanafir Island, 4.U.49. 

The transverse streaks characteristic of the male and the pale areas 
overlying the iridescent patches are clearly shown in the photograph. 

PLATE 29 

Figs. 3 and 4. Sepioteuthis sp. ; dorsal and ventral views of a young 
immature specimen taken off Sherm Sheik, i.ii.49. 

Fig. 5. Sepioteuthis lessoniana Lesson; left tentacle club of $ shown on 
Plate 28, fig. 1. 

Fig. 6. Sepioteuthis lessoniana Lesson ; right tentacle club of $ taken at 
Aqaba, 28.1.49. 

Fig. 7. Octopus horridus Orbigny taken at Tiran Island, io.i.49. 

PLATE 30. OCTOPUS CYANEA GRAY 

Fig. 8. Lateral view of a $ taken at Sherm Sheik, 12.1.49. 

Fig. 9. Oral face of the same specimen as in Fig. 8. The so-called ' zebra ' 
markings on the lateral side of the arms are a constant feature in this 
species. 



Bull. B.M. {N.H.) Zoology, I. 8 



PLATE 28 




& 



SEPIOTEUTHIS LESSONIANA LESSON 



Bull. B.M. {N.H.) Zoology, I, 8 



PLATE 29 










AQABA CEPHALOPODA 



Bull. B.M. (N.H.) Zoology, I, 8 



PLATE 30 











OCTOPUS CYANEA GRAY 



VII. ECHINODERMATA 

By AILSA M. CLARK 

The collection of Echinoderms includes many well-known littoral species which are 
widespread throughout the Indo-West Pacific area, as well as some which are peculiar 
to the Red Sea. A few species, notably the single Crinoid Capillaster multiradiata 
(Linnaeus) and an Echinoid, Clypeaster fervens Koehler, have not been previously 
recorded from the Red Sea. 

The species are the following, all of them from low tide or low spring tide level 
except where otherwise stated. Those mentioned in more detail in the text are 
marked with an asterisk. References in the text giving further details are marked 
with a dagger. 



ASTEROIDEA 



Locality 



Number 



Astropecten poly acanthus Miiller & Troschel 


Dahab 


i 




Ras Muhammad Bay 


i 


*Fromia ghardaqana Mortensen 


Dahab 


i 




Abu Zabad 


3 


*Gomophia egyptiaca Gray 


Abu Zabad 


i 


Linckia multifora (Lamarck) . 


Sherm Sheik 


2 




Dahab 


I 




Sanafir I. 


I 


Aster ope carinifera (Lamarck) 


Abu Zabad 


2 


*Asterina burtonii Gray .... 


Sanafir I. 


I 




Dahab 


I 




Sherm Sheik 


I 




Abu Zabad 


4 


OPHIUROIDEA 






*Ophiocoma pica Miiller & Troschel . 


. Dahab 


5 




Sherm Sheik 


4 




Sanafir I. 


5 




Mualla 


i 




Tiran 


3 




Abu Zabad 


5 


*Ophiocoma scolopendrina (Lamarck) 


Sanafir I. 


3 




Dahab 


8 




Faraun Id. 


IO 




Sherm Sheik 


4 




Abu Zabad 


2 


*Ophiocoma erinaceus Miiller & Troschel . 


Dahab 


2 




Sherm Sheik 


I 




Sanafir I. 


2 




Abu Zabad 


2 


Ophiocoma valenciae Miiller & Troschel . 


Tiran 


4 




Sanafir I. 


i 




Sherm Sheik 


i 




Abu Zabad 


2 




Dahab 


I 


*Ophiocoma sp. ..... 


Sherm Sheik 


I 


*Macrophiothrix hirsuta (Miiller & Troschel) 


Sherm Sheik 


I 




Sanafir I. 


I 




Dahab 


2 



204 THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 

Locality Number 

Ophiotrichoides propinqua (Lyman) . . Dahab 2 

* Placophiothrix purpurea (von Martens) . . Dahab 1 

Ophiolepis cincta Muller & Troschel . . Dahab 5 

Abu Zabad 2 

Sherm Sheik 3 



CRINOIDEA 

'Capillaster multiradiata (Linnaeus) . 



Dahab 



ECHINOIDEA 

Eucidaris metularia (Lamarck) 



Diadema setosum (Leske) 



Echinometra mathaei (Blainville) 



Heterocentrotus mammillatus (Linnaeus) 
*Tripneustes gratilla (Linnaeus) 



Clypeaster humilis (Leske) 
*Ciypeaster fervens Koehler 
Lovenia elongata (Gray) 



Sherm-el-Moiya 

Mualla 

Sherm Sheik 

Sanafir I. 

Tiran 

Aqaba 

Sherm-el-Moiya 

Tiran 

Faraun I. 

Abu Zabad 

Mualla 

Abu Zabad 

Sherm Sheik 

Tiran 

Sanafir I. 

Dahab 

Dahab 

Abu Zabad 

Sanafir I. 

Dahab 

Aqaba 

Dahab 

Dahab 

Dahab 



4 
4 
5 
4 
10 

3 
3 

1 
6 

2 
1 
1 
1 



HOLOTHUROIDEA 

Synapta maculata (Chamisso & Eysenhardt) 



Synaptula recta (Semper) 
Halodeima edulis (Lesson) 

Halodeima atra (Jager) .... 
Halodeima cinerascens (Brandt) 
Holothuria impatiens (Forskal) 
*Holothuria sucosa Erwe. 
Holothuria pardalis (Selenka) . 

Holothuria curiosa var. pervicax (Selenka) 
Microthele difficilis (Semper) . 

*Microthele nobilis (Selenka) . 
Actinopyga miliaris (Quoy & Gaimard) . 



Um Nageila (in 


shallow 


I 


water off mangrove 




swamp) 






Sherm Sheik 




1 (pt.) 


. Dahab 




1 


Abu Zabad 




1 


Abu Zabad 




2 


Abu Zabad 




1 


Dahab 




5 


. Dahab 




1 


. Dahab 




5 


Graa 




2 


Dahab 




1 


Abu Zabad 




8 


Dahab 




6 


Ras Muhammad Bay 


1 


Faraun I. 




1 



THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 205 

ASTEROIDEA 

Family Linckiidae 

Fromia ghardaqana Mortensen 

Pl. 31, Figs, a-c 

Scy taster milleporellus, Miiller and Troschel, 1842 : 35 ; [non Asterias milleporella Lamarck, 18 16: 

564]. 
Fromia milleporella (part), Gray, 1866: 14. 
Fromia monilis, Tortonese, 1935: 70; 1936: 213; [non Fromia monilis Perrier, 1875: 443 (p. 179 

in repaged edition)]. 
Fromia ghardaqana Mortensen, 1938: 37. 

Dahab, shore ; 1 specimen. Abu Zabad, reef at low water springs ; 3 specimens. 

Description. R = 40 mm., r = 10 mm., R/r = 4-0. The arms taper evenly 
throughout their length to a rather pointed tip. One has been broken and is in pro- 
cess of regeneration. Of the five primary inter-radial plates, three are enlarged with a 
flat surface raised slightly above the level of the surrounding plates, the one adjacent 
to the madreporite is smaller but also a little elevated, while the fifth is not at all 
conspicuous. The madreporite is triangular in shape, with deep radiating grooves, 
and measures 1-4 mm. across. 

The carinal row of plates is not very clear proximally, where all the plates are in 
fact rather irregularly arranged. At the base of the arm there are about seven plates 
across the width. 

All the dorsal and ventral plates, as well as the marginals, are closely covered with 
uniform, smooth, rounded granules, about 7 in the length of 1 millimetre. These lie 
very close together and are polygonal on the convex plates, of which there are about 
10 on the dorsal side of each arm, besides the primary inter-radial plates and the 
marginals. Most of the convex plates are near the tip of the arm, but an irregular 
series of spaced plates occupies the mid-radial distal area. 

The number of supero-marginal plates varies between 19 and 21 on the four com- 
plete arms, with the same number in each infero-marginal series. The latter plates 
are relatively narrower and are noticeably longer than broad. In the distal half of 
the arm they may bear a small tubercle in the centre as also do the last two supero- 
marginals. These plates, unlike the infero-marginals, are not evenly sized but, especially 
distally, large and small plates tend to alternate with one another, the larger ones 
being rather convex. The two series of marginal plates tend to alternate in position. 

On the ventral side the papulae are clearly visible in the angles between the plates. 
The granules surrounding each one are not markedly larger than the other granules. 
Proximally there are 3 rows of papulae, correlated with the presence of 4 rows of 
ventro-lateral plates. The outermost row of these consists of only 4 plates on each 
side of the interbrachial angle, extending to the aboral end of the second infero- 
marginal plate. The third series reaches the seventh infero-marginal, the second to 
the eleventh plate, and the innermost series to the fourteenth. 

The adambulacral plates bear 2, or, in the middle part of the arm, often 3 flattened 
furrow spines. Outside these is a single stumpy spine, shorter than the furrow spines 
though much thicker and slightly elongated in transverse section. On both sides of 
this spine and outside it are numerous granules like those of the ventro-lateral plates. 



206 THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 

Remarks. Muller and Troschel's description of Scytaster milleporellus together with 
the locality of the Red Sea suggests that their specimens like many of Gray's were 
almost certainly Fromia ghardaqana. However, some of the latter, from Mauritius 
and other localities in the Indian Ocean, are the form (pi. 31, fig. d) with even-sized 
supero-marginal plates which is generally assumed to represent Fromia milleporella. 
Since Lamarck gave as the type locality 'les mers d'Europe?', and only a brief 
description, it is not certain which form really is milleporella. This question can 
only be answered by study of the type specimens if they are still in existence. 

Mortensen has examined the type of Fromia monilis Perrier and finds it quite 
distinct from the Red Sea species, although he does not give any details. 

On comparison of the specimen described with one of F. monilis from Macclesfield 
Bank, South China Sea, with R = 35 mm., it is at once seen that the granulation of 
the dorsal side of the latter is very much finer with at least 10 granules to a millimetre 
rather than only 7. Also the arms of F. monilis are relatively much narrower with 
R/r about 4-5 on average and the supero-marginal plates usually occupy more of the 
dorsal surface of the arm, so that only 3 or 5 rows of plates, rarely more, lie across 
the base of the arm. On the ventral side the granules around the pores are clearly 
enlarged unlike those of F. ghardaqana. 

Unfortunately there are no specimens of Fromia pacifica H. L. Clark (the species 
that Mortensen says is most nearly related to F. ghardaqana) in the British Museum to 
compare with the material from the Red Sea. That Torres Strait species apparently 
has even-sized supero-marginal plates and pointed granules rather than flat ones. 

There are also three juvenile specimens in the present collection, the two larger 
ones having R = 18 mm., but whereas one is much more slender with an R/r ratio 
of 3*5 :i, the other has the ratio only 2-8. Of the many old dry specimens in the British 
Museum, the R/r value varies between 3-0 and 37, although a co-type of F. gharda- 
qana from Ghardaqa sent by Dr. Mortensen has the ratio 4-0. This it seems is just 
about the maximum value. 

From all the other species of Fromia, F. ghardaqana is easily distinguished by the 
alternate large and small distal supero-marginals. 



Gomophia egyptiaca Gray 
Pl. 32 

Gomophia egyptiaca Gray, 1840: 286. H. L. Clark, 1921: 55. 

f Scytaster aegyptiacus, Perrier, 1875: 428 (p. 164 in re-paged edition). 

Nardoa aegyptiaca, de Loriol, 1891: 30. Fisher, 1906: 1087. Koehler, 1910: 157, pl. xvii. 5, 6. 

Abu Zabad, reef at low water springs ; one specimen. 

R = 84 mm., while the type has R = 62 mm. The intermarginal plates in the arm 
angle are not more conspicuous than in the type and indeed are quite hidden by the 
granulation in one of the angles. 

Range. Red Sea, Mauritius, Samoa, Philippines, Fiji, Macclesfield Bank. 



THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 207 

Family Asterinidae 

Asterina burtonii Gray 

Asterina burtonii Gray, 1840: 289. fG. A. Smith, 1927: 641. 

Asteriscus wega Perrier, 1869: 102. 

Asterina wega Perrier, 1876: 238 (p. 318 of re-paged edition). 

Sanafir ; one specimen. Dahab ; one 6-armed specimen. Abu Zabad ; 4 specimens. 
Sherm Sheik ; one 7-armed specimen. 

Remarks. Since in 1876 Perrier corrected the error in his original description of 
A. wega, regarding the number of spines on each ventro-lateral plate, emending it to 
2 or 3 rather than 1, there seems to be no reason why specimens with up to 8 arms 
should not be regarded as Asterina burtonii. Smith accepts 6-armed specimens as 
such. These forms with more than 5 arms are usually juvenile and more or less 
obviously in process of regeneration. The 7-armed specimen in the present collection 
has 4 arms diminutive. Perrier states that all his thirteen specimens of A . wega were 
undergoing regeneration. 

2. OPHIUROIDEA 
Family Ophiocomidae 

Ophiocoma pica Muller & Troschel 

Ophiocoma pica Muller & Troschel, 1842: 101. H. L. Clark, 1921: 127, pi. xiii, 8 (coloured). 

fEly 1942: 54, pi. xii, B.i., text-fig. 15. 
Ophiocoma lineolata Muller & Troschel, 1842: 102. de Loriol, 1893: 28. 

Dahab ; 5 specimens. Sherm Sheik ; 4 specimens. Sanafir ; 5 specimens. Mualla ; 
1 specimen. Tiran ; 3 specimens. Abu Zabad ; 5 specimens. All from coral at low tide. 

Remarks. These specimens are easily distinguished from the other Ophiocomas 
collected by the conspicuous stripes on the otherwise black arms and the yellowish 
stripes on the disk. The ratio of arm length to the disk diameter varies between 
3-6 and 4-8: 1. 

Note. It has been accepted for a very long time that 0. pica and 0. lineolata are 
synonymous, but both names have been retained by different authors. For instance 
Koehler (1922a; : 324) still uses lineolata although most other recent authors prefer pica. 
However, the latter name had page priority in Muller & Troschel's System der 
Asteriden. So in spite of its previous use in manuscript by Valenciennes, which has 
no validity, the name Ophiocoma lineolata should be dropped. 

Ophiocoma scolopendrina (Lamarck) 

Ophiura scolopendrina Lamarck, 1816, 2: 544. 

^Ophiocoma scolopendrina, de Loriol, 1893: 23. H. L. Clark, 1921: 125, pi. xiii. 9. f Koehler, 
1922a: 325, pis. lxxiii. 5; lxxiv. 1-7. 

Sanafir ; 3 specimens. Dahab ; 8 specimens. Faraun Island ; 10 specimens. Sherm 
Sheik ; 4 specimens. Abu Zabad ; 2 specimens. All from the shore under stones, 
zoo. 1. 8. Ee 



208 THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 

Remarks. The colour ranges from variegated bluish grey to dense black on the 
dorsal side of the disk and arms, the ventral side of the disk being always pale. Most 
have the arms broken but they are usually relatively long, six or more times the disk 
diameter. 

Ophiocoma erinaceus Muller & Troschel 

Ophiocoma erinaceus Muller & Troschel, 1842: 98. fde Loriol, 1893: 21. H. L. Clark, 1921: 127. 
jEly, 1942: 52, text-fig. 45, pi. xiia. 

Dahab: 2 specimens. Sherm Sheik; 1 specimen. Sanafir; 2 specimens. Abu 
Zabad: 2 specimens. 

Remarks. Except for the two specimens from Abu Zabad, these are densely black 
all over ; even the tentacles of those from Dahab are black ; also the arms are relatively 
short, the ratio of arm length to disk diameter being 4-4-8 : 1. The Abu Zabad speci- 
mens are also densely black dorsally but are pale on the underside of the disk, 
although the tentacles are black. The arms of one are all broken but in the other 
their length is nearly seven times the disk diameter. They are thus intermediate 
between 0. scohpendrina (with relatively long arms and lighter colour) and 0. 
erinaceus, with shorter arms and a uniformly dark colour, so there was some doubt 
as to which species they should be. Finally they were referred to the latter species 
for the following reasons : besides the very dense black colour on the dorsal side, the 
disk granulation hardly extends below the periphery and there are two tentacle scales 
for quite a large part of the arm, as in erinaceus. Also, apart from these morphologi- 
cal characters, the fact that they were taken well out on the reef at low spring tide 
level in the same zone as Ophiocoma pica suggests that they belong to erinaceus, 
for H. L. Clark makes the distinction of habitat of the two forms scohpendrina and 
erinaceus an important reason for maintaining them as separate species, the former 
characteristically occupying a higher level on the shore which is uncovered at ordinary 
low tides. 

I fully agree with Ely that very rarely can several characters be used to distinguish 
intermediate specimens as belonging to one or the other species. Quite often con- 
flicting results are obtained by using two different characters. For instance there is a 
specimen in the British Museum collection from Muscat, with the proportions 
170 mm./2i mm. = 6-4:1, which would on this count be called scohpendrina, but the 
unrelievedly black colour on the contrary suggests that it is erinaceus. In such cases 
only a detailed observation of the habit and habitat can produce a conclusive 
identification. 

Ophiocoma sp. 

Sherm Sheik; 1 specimen. 

This is a very small specimen (disk diameter = 5 mm.) with all the arms broken 
and a hole through the centre of the disk. It is nearest to O.pica as there are 2 tentacle 
scales, 5 slender arm spines proximally, and dark bands on the arms, also the oral 
shields are longer than wide. However, the dorsal side of the disk is unusual in having 
black spots each surrounded by a lighter ring on a dark brown background. These 
spots vary in size and shape but are relatively much larger than those of Ophiocoma 
doderleini. 



THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 209 

Family Ophiotrichidae 

Placophiothrix purpurea (von Martens) 

Ophiothrix purpurea von Martens, 1867: 346. Doderlein, 1896: 296, pis. xiv. 12; xvii. 23. 
^Ophiothrix lepidus de Loriol, 1893: 45, pi. xxv. 1. 
f Ophiothrix fallax de Loriol, 1893: 47, pi. xxv. 2. 
Placophiothrix purpurea, H. L. Clark, 1939: 86. 

Dahab ; i specimen. 

This specimen agrees very closely with de Loriors description of Ophiothrix fallax 
from Mauritius, as it has a pale green disk and relatively long arms (disk diameter 
= 4*5 mm., arm length = 45 mm.). H. L. Clark has declared 0. lepida de Loriol to 
be a synonym of 0. purpurea, from a study of the long series of specimens obtained 
by the John Murray Expedition. He makes no mention of 0. fallax, but as the 
characters of that species are intermediate between those of the other two, it cer- 
tainly comes within the range of variation of Placophiothrix purpurea. 

Possibly Doderlein's Ophiothrix lorioli (1896: 297) from Amboina, with radial 
shields similar to those of 0. lepida, is also a synonym of purpurea. Both Doderlein 
and Koehler (1898: 102) say that 0. lepida and 0. lorioli cannot be confounded, but 
neither of them give any reason for this. 



Macrophiothrix hirsuta (Miiller & Troschel) 

Ophiothrix hirsuta Miiller & Troschel, 1 842 : 1 1 1 . Marktanner-Turneretscher, 1887:311. f Koehler, 

1922a: 234, pis. xxxi. 1, 2; xxxiii. 13; xcix. 2. Tortonese, 1949: 37. 
Ophiothrix cheneyi Lyman, 1861: 84. 
Macrophiothrix hirsuta, H. L. Clark, 1938: 285. 
Ophiothrix demessa, H. L. Clark, 1939: 83. [non Ophiothrix demessa Lyman, 1861: 82.] 

Sherm Sheik ; 1 specimen. Sanafir ; 1 specimen. Dahab ; 2 specimens. 

Remarks. There seems to be considerable difference of opinion as to the shape of the 
dorsal arm-plates in this species. H. L. Clark describes them as more or less oval in 
his key to the species of Macrophiothrix, but as Tortonese points out, Miiller & 
Troschel's original description mentions lateral angles, a statement open to several 
interpretations but suggesting at least something a little more angular than an 
ellipse. Koehler's plate 83, fig. 13, of the arm of a Philippine specimen shows dorsal 
arm-plates of which the widest part is midway between proximal and distal edges, 
whereas all the Red Sea specimens that I have seen have the widest part distinctly 
distal to the half-way line with a slightly rounded angle as opposed to the very acute 
angle of M. longipeda. This rather fan-shaped form is shown in Koehler's plate 31, 
fig. 1, of a specimen from the Red Sea, which also resembles the present material in 
the characters of the disk. That the shape of the dorsal arm-plates varies in different 
parts of the range is shown by the fact that Lyman's species from Zanzibar, 0. 
cheneyi, which is commonly accepted as a synonym of M. hirsuta, is described as 
having oval, microscopically granulated dorsal arm-plates. 



210 THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 

The latter feature, that is the presence of more or less thorny granules on the 
dorsal arm-plates, is not mentioned by Miiller & Troschel, but Marktanner-Turner- 
etscher states that they are always somewhat granulated although this is not so 
marked as in 0. demessa. In fact he considers the difference in the size and thorni- 
ness of these granules to be the only difference separating the two species. Through 
the courtesy of Dr. Elisabeth Deichmann I have had the opportunity of studying 
some specimens of 0. demessa and as a result fully agree with Marktanner-Turner- 
etscher, the only other difference that I can see being that the arms seem to taper 
more rapidly, in younger specimens at least, of 0. demessa. The granules on the arms 
are distinctly more thorny than in the specimens from the Red Sea, where they may 
be quite unobtrusive in spirit specimens. H. L. Clark in his John Murray Report 
names two specimens from the Red Sea and the Gulf of Aden Ophiothrix demessa, 
of which the one in the British Museum is indistinguishable from M. hirsuta, and I 
suspect that Koehler's record of 0. demessa from the Red Sea is also based on a 
similar specimen. In 1946 H. L. Clark erected a new genus Amphiophiothrix to 
accommodate the species 0. demessa, but I cannot agree that there is a generic dis- 
tinction between it and Macr ophiothrix hirsuta. 

The validity of some of the other Indo-Pacific species of Macr ophiothrix has been 
questioned by several authorities. Some of them are possibly variants of other 
species such as hirsuta in which the granulation of the radial shields is reduced, for 
there is a tendency for such a reduction throughout the genus as there is also for the 
development of granules on the dorsal arm-plates, a character featuring in the 
descriptions of several species, such as M. rugosa H. L. Clark, and noticeable also in 
some larger specimens of other species. However, without seeing the types and being 
able to compare them with large series of specimens from different parts of the Indo- 
Pacific, it is impossible to add anything concrete to the suspicions already voiced. 



3. CRINOIDEA 
Family Comasteridae 

Capillaster multiradiata (Linnaeus) 

Asterias multiradiata, Linnaeus, 1758: 663. 
Capillaster multiradiata, A. H. Clark, 1909: 364. 

^Capillaster multiradiata, A. H. Clark, 1931: 173, pis. iii. 5; xi. 30; xiii. 34; xiv. 35, 36; lxxxi. 
222, 223, also many text-figs. 

Dahab ; 1 specimen ; arms 90 mm. in length. 

This is the first record of this species from the Red Sea, the former known range 
being from Formosa south to northern Australia and west as far as the Maldive 
Islands, so its discovery here is most interesting. 

There are 36 arms, which is rather more than usual ; A. H. Clark gives 12 to 35 
as the usual range, but quotes specimens with up to 43 arms. 



THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 211 

4. ECHINOIDEA 
Family Toxopneustidae 

Tripneustes gratilla (Linnaeus) 

Echinus gratilla Linnaeus, 1758: 664. 

Tripneustes gratilla, Loven, 1887: 77. fMortensen, J943. 3 (2) : 500, pis. xxxiii. 1-3; xxxiv. 2-6; 
xxxv. 3-4; xxxvii. 1-2, 4-10; xxxviii. 1-4; lvi. II. 

Abu Zabad, reef at low spring tide ; 3 specimens. Sanafir ; 1 specimen. Dahab ; 
6 specimens. Aqaba ; 2 specimens. 

The two from Aqaba are superficially very different from the others, having 
relatively few and long primary spines above the ambitus, which are white in colour 
and contrast sharply with the dark purple of the test, produced mainly by the numer- 
ous pedicellariae. The tube feet of these two specimens are black or at least have a 
black band around them. The other specimens are more drab in colour, several being 
slightly reddish and their tube feet are grey. The denuded tests are distinctly green 
aborally. 

Family Clypeastridae 

Clypeaster (Rhaphidoclypus) fervens Koehler 

Clypeaster fervens Koehler, 1922: 45, pis. vi. 1, 2, 6; xv. 1. 

^Clypeaster {Rhaphidoclypus) fervens, Mortensen, 1948, 4 (2): 84, pis. xiii. 2, 3; xxii. 1-11; 
xxvi. 2; lxv. 7-9, 12, 20. 

Dahab, shore ; 1 dead test. 

This specimen is easily distinguished from Clypeaster humilis by the relatively 
large petals and the concave oral side. It is 46 mm. in length but already has well- 
developed genital pores. According to Dr. Mortensen (who has very kindly confirmed 
my identification) in his monograph, the genital pores only begin to appear when the 
length is about 56 mm., that is in the John Murray Expedition material from the 
Indian Ocean. It seems then that in the Red Sea this species undergoes precocious 
genital development. 

5. HOLOTHUROIDEA 

Family Holothuriidae 

Holothuria sucosa Erwe 

Cucumaria hartmeyeri Heifer, 1912 : 332. [non Holothuria hartmeyeri Erwe, 1913 : 383, pi. vii. 19.] 
^Holothuria sucosa Erwe, 1919: 186, text-fig. 5. Panning, 1934, 3: 80, text-fig. 64. 
? Holothuria ocellata, Tortonese, 1936: 235, text-figs. 5, 6. 

Dahab ; 1 specimen. 

The knobbed buttons have 4 or 5 pairs of holes, sometimes as many as 10 pairs. 
Unlike H. arenicola var. boutani Herouard, which also has multilocular, though flat 
buttons, the tables, which are also larger, have a complete ring of holes around the 
margin not interrupted by the extended four central holes. Unlike H. ocellata 
Jager, the great majority of buttons have more than 3 pairs of holes. 



212 THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 

Microthele nobilis (Selenka) 

Mulleria nobilis Selenka, 1867: 31, pi. xvii. 13-15. 

\Holothuria {Microthele) nobilis, Panning, 1929, 1: 131, text-fig. 15. 

Microthele nobilis, Heding, 1940: 320. 

Ras Muhammad ; 1 specimen. 

Although shrunken in preservation this specimen still measures 24 cm. in length. 
The tables have mostly rather irregular disks. The other dorsal deposits are 'three- 
dimensional buttons ', fenestrated irregularly with about 4 pairs of holes on each face. 
Ventrally, however, these spicules are much outnumbered by more conventional 
flat buttons with holes in one plane, there being usually 4 or 5 pairs of holes if not 
more. 

REFERENCES 

Clark, A. H. 1909. The Crinoids of the 'Gazelle' Expedition. Zool. Anz. 34 (11-12): 363-376. 
1931. A monograph of the Existing Crinoids. 1 (3). Bull. U.S. Nat. Mus. 82: vii, 1-816, 

82 pis. 
Clark, H. L. 1921. The Echinoderm fauna of Torres Strait. Pap. Tortugas Lab. 10: vi, 1-223, 

38 pis. 
1938. Echinoderms from Australia. Mem. Mus. comp. Zool. Harv. 55: viii, 1-596, 

text-figs. 1-64, pis. 1-24. 

1939. Ophiuroidea. Sci. Rep. John Murray Exped. 6: 29-136, text-figs. 1-62. 

1946. The Echinoderm fauna of Australia. Pub. Carneg. Instn. 566: iv, 1-567. 

Doderlein, L. 1896. Bericht fiber die von Herrn Prof. Semon bei Amboina und Thursday- 
Island gesammelten Ophiuroidea. In Semon, Zoologische Forschungsreisen in Australien 

und dem Malayischen Archipel. pp. 279-300, pis. 14-17. Jena. 
Ely, C. A. 1942. Shallow water Asteroidea and Ophiuroidea of Hawaii. Bull. Bishop Mus. 

Honolulu, 176: 1-63, text-figs. 1-18, pis. 1-13. 
Erwe, W. 1913. Holothuroidea. In Michaelsen, W., & Hartmeyer, R., Die Fauna Sudwest- 

Australiens. Ergebn. Hamburg, siidwest-austr. Forsch. 4: 349-402, text-fig. 1, pis. 5-8. 

1919. Holothurien aus dem Roten Meer. Mitt. Zool. Mus. Berlin, 9 (2): 177-189. 

Fisher, W. K. 1906. The Starfishes of the Hawaiian Islands. Bull. U.S. Fish. Comm. 1903. 

3: 987-1130, pis. 1-49. 
Gray, J. E. 1840. A Synopsis of the genera and species of the class Hypostoma. (Asterias 

Linn.) Ann. Mag. Nat. Hist. 6: 175-184, 275-290. 
1866. Synopsis of the Species of Starfish in the British Museum, iv-f 17 pp., pis. 1-16. 

London. 
Heding, S. G. 1940. Die Holothurien der Deutschen Tiefsee Exped. II. Aspidochirote und 

Elasipode Formen. Wiss. Ergebn. Valdivia, 24: 317-375, text-figs. 1-2 1. 
Helfer, H. 191 2. tJber einige von Dr. Hartmeyer im Golf von Suez gesammelte Holothurien. 

Mitt. Zool. Mus. Berlin, 6 (2) : 327-334. 
Koehler, R. 1898. fichinodermes recueillis par 1" Investigator' dans l'Oc6an indien. II. Les 

ophiures littorales. Bull. Sci. France Belgique, 31: 55-124, pis. 2-4. 

1910. Echinoderma of the Indian Museum. 6. Asteroidea II. pp. 1-191, pis. 1-20. Calcutta. 

1922. Echinoderma of the Indian Museum. 8. Echinoidea. II. Clypeastrides et Cassidulides. 

pp. 1-161, pis. 1-15. Calcutta. 
1922a. Ophiurans of the Philippine Seas. Bull. U.S. Nat. Mus. 100 (3). x, 1-486, pis. 

1-103. 
Lamarck, J. B. P. A. de Monet de. 181 6. Histoire naturelle des animaux sans vertebres. II. 

pp. 1-568. Paris. 
Linnaeus, C. 1758. Sy sterna Naturae. Ed. X. 1. 824 pp. Holmiae. 



THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 213 

Loriol, P. de. 1 891. Notes pour servir a l'etude des Flchinodermes. III. Mem. Soc. Phys. 

Geneve. Suppl. 1890 (8): 1-31, pis. 1-3. 
1893. Catalogue raisonne des iEchinodermes recueillis. par M. V. de Robillard a l'He 

Maurice. III. Ophiurides et Astrophy tides. Mem. Soc Phys. Geneve, 32 (1): 1-59, pis. 

23-25- 
Loven, S. 1887. On the species of Echinoidea described by Linnaeus in his work 'Museum 

Ludovicae Ulricae'. Bih. Svensk Vetensk. Akad. Handl. (13) 4 (5): 1-185, pis. 1-9. 
Lyman, T. 1861. Descriptions of new Ophiuridae. Proc. Boston Soc. Nat. Hist. 8: 75-86. 
Marktanner-Turneretscher, G. 1887. Beschreibung neuer Ophiuriden und Bemerkungen 

zu bekannten. Ann. naturh. (Mus.) Hofmus. Wien, 2: 291-316, pis. 12-13. 
Martens, E. von. 1867. Uber vier neue Schlangensterne. Mber. preuss. Akad. wiss.: 345-348. 
Mortensen, Th. 1926. Cambridge Expedition to the Suez Canal in 1924. VI. Echinoderms. 

Trans. Zool. Soc. Lond. 22: 117-131, text-figs. 11-13. 
• 1928-1948. Monograph of the Echinoidea. Copenhagen. 1. Cidaroidea, 1928; 3 (1) Aulo- 

donta, 1940; 3 (2) and (3) Camarodonta, 1943; 4 (2) Clypeastroida, 1948. 
1938. Contributions to the study of the Development and Larval forms of Echinoderms. 

IV. K. Danske Vidensk. Selsk. Skr. (9) 7 (3) : 1-59, text-figs. 1-30, pis. 1-12. 
Muller, J., & Troschel, F. H. 1842. System der Asteriden. xx + 134 pp., pis. 1-12. Braun- 
schweig. 
Panning, A. 1929-1935. Die Gattung Holothuria. Parts I-V. Mitt. Zool. St. Inst. Hamburg, 

44: 91-138, text-figs. 1-21 ; 45: 24-50, 65-84, 85-107, text-figs. 22-102; 46: 1-18, text-figs. 

103-121. 
Perrier, E. 1869. Recherches sur les pedicellaires et les ambulacres des Asteries et des Oursins. 

I. Ann. Sci. nat. (5) 12: 197-304, pis. 17, 18. 
1875. Revision de la collection de Stellerides du Museum d'Histoire Naturelle de Paris. 

384 pp. Paris. (Also published in Arch. Zool. exp. gen. 4 (1875) : 263-449 ; 5 (1876) : 1-104, 

209-304). 
Selenka, E. 1867. Beitrage zur Anatomie und Systematik der Holothurien. Z. wiss. Zool. 

17: 291-374, pis. 17-20. 
Smith, G. A. 1927. On Asterina burtonii Gray. Ann. Mag. Nat. Hist. (9) 19: 641-645. 
Tortonese, E. 1935. Gli Echinodermi del Museo di Torino. III. Asteroidi. Boll. Mus. Zool. 

Anat. comp. Torino, 45 (3): 27-132, pis. 1-11. 

1936. Echinodermi del Mar Rosso. Ann. Mus. Stor. nat. Genova, 59: 202-245, text-figs. 1-8. 

1949. Echinodermi della Somalia Italiana. Ann. Mus. Stor. nat. Genova, 64: 30-42, 1 pi. 



Legends to Plates 31 and 32 

PLATE 31 

Fromia ghardaqana Mortensen, specimen from Dahab. (a) Dorsal side ; 
(b) ventral side; (c) specimen 40.3.23.35; and (d) Fromia milleporella 
Lamarck, specimen 39.3.29.20, for comparison. 

PLATE 32 

Gomophia egyptiaca Gray, (a) Dorsal side of the type and (b) an inter- 
brachial angle to show the intermarginal plates. 



Bull. B.M. (N.H.) Zoology, I, 8 



PLATE 31 




5^*^ 



FR0M1A GHARDAQANA MORTENSEN (Figs, a -c) 
FROMIA MILLEPORELLA LAMARCK (Fig. d) 



Bull. B.M. {N.H.) Zoology, I, 8 



PLATE 32 




GOMOPHIA EGYPTIACA GRAY 



VIII. TUNICATA 

By WILLARD G. VAN NAME 

AMERICAN MUSEUM OF NATURAL HISTORY 

Through the kindness of the authorities of the British Museum (Natural History) 
the Tunicata collected by the M. Y. Manihine in the Gulf of Aqaba in the early months 
of 1949 were forwarded to me for examination. 

As far as I am aware no collection of Tunicata has previously been made there, but 
the tunicates of the Red Sea, and especially those of the Gulf of Suez, have been the 
subject of much study and are dealt with in several published articles. 

The remarkable work by Savigny (18 16), which was many decades in advance of 
his time, and which laid the foundations of much of our knowledge of the Tunicata, 
as well as important articles by Hartmeyer, Michaelsen, and others during the present 
century, were based in large part on specimens from those waters. 

It was therefore hardly to be expected that new species would be found in a com- 
paratively small collection, especially since no specimens were obtained except in 
very shallow water, in no case over about 2 fathoms. 

All the specimens appear to be referable to species already described, but never- 
theless the collection contains some that are of interest, especially those of the solitary 
form of Salpa maxima var. tuber culata described by Metcalf, 191 8, from the southern 
Philippines, who, however, had specimens of the aggregated form only. 

Since the Gulfs of Suez and Aqaba are extensions of the Red Sea and consequently 
of the tropical part of the Indian Ocean, their faunas are Indo-Malayan, in spite of 
their near approach geographically to the eastern Mediterranean. 

This fact is, however, not so evident in the present collection as might be expected, 
since it happens to contain some species that are practically circumtropical, and found 
both in the Mediterranean and Indian Ocean. These species are shallow water forms, 
and it is possible that some of them may owe their very extensive distribution to 
human agency, by transportation on the bottom of ships. 

The Tunicata in this collection appear to belong to the following 13 species, one of 
them (Salpa maxima) being perhaps represented by two varieties: 

Class ASCIDIACEA 
Compound Ascidians 

1. Polyclinum saturnium Savigny, 1816 
Polyclinum saturnium Savigny, 1816: 190, pi. 19, fig. 1; Michaelsen, 1920: 9. 

One rather thick colony measuring over 50 mm. in extent. 

2. Didemnum candidum Savigny, 1816 

Didemnum candidum Savigny, 1816: 194, pi. 4, fig. 3, pi. 20, fig. 1. 

Several small colonies with abundant spicules, whose points are so short and 
zoo. 1. 8. Ff 



216 THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 

slightly developed that the spicules are almost spherical. Also one small colony 
having spicules with larger and better developed rays or points. 

There is also one colony, growing on coral, which has very few spicules and a great 
many faecal pellets in the intestinal tracts of the zooids, perhaps indicating an 
incipient case of the so-called ' Hypurgon ' condition to which this and allied forms 
are subject, in which the water currents in the cloacal canals become too weak to 
carry off the waste material, which remains in the cloacal system and in the common 
test, greatly altering the character and appearance of the colony, but there does not 
seem to be any reason for assuming that it is of a different species. See Michaelsen, 
19190: 11-13. 

Didemnum candidum appears to be a species of very wide distribution, being found 
also in American waters, very abundantly in some places. 

It cannot be doubted that far too many species of the genus Didemnum have been 
described. Apparently this is in part due to overlooking the great effects on the 
general appearance of the colony of its age and past history, particularly in the case 
of old colonies. Many or most of the species are subject to periods (in many cases 
seasonal) of regression and extensive degeneration of the zooids, followed by sub- 
sequent recovery and regrowth of the colony to its normal functional condition. 
During such regressive periods, though the zooids degenerate more or less com- 
pletely, the spicules may endure unchanged through several or perhaps many genera- 
tions of the zooids. The result is that in old colonies we may find a far greater 
abundance of spicules than the spicule-forming ability of the zooids present could 
possibly account for, and likewise often peculiarities in the distribution of the 
spicules, which one must not mistake for specific characters. Old colonies are apt to 
acquire a hard calcareous character in which the spicules form a far larger component 
than the test substance and zooids do. 

Simple Ascidians 

3. Phallusia nigra Savigny, 1816 

Phallusia nigra Savigny, 1816: 163, pi. 2, fig. 2; pi. 9, fig. 1. 

Ascidia atra Lesueur, 1823: 2, pi. 1, fig. 2. 

Ascidia nigra, Herdman, 1882: 210. 

Phallusia nigra, Hartmeyer, 1916: 408, figs. 5-9. 

Eleven specimens, all of small size. This species, widely distributed and common 
in shallow water in many warm regions of both hemispheres, is easily recognizable 
from its bluish or blue-black coloration. 

If Phallusia is accepted as a genus distinct from Ascidia, the present species should 
be placed in it, as in old and large individuals the neural duct has accessory apertures, 
at least in many specimens. In other respects it is a very typical Ascidia. 

4. Phallusia sp., apparently Phallusia arabica Savigny, 1816 
Phallusia arabica, Hartmeyer, 1916: 414, figs. 10-12. 

One specimen of 52 mm. body length (or 63 mm. if the obliquely forwardly ex- 
tending atrial siphon is included). In external features other than unusual forward 



THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 217 

position of the atrial siphon (probably only an individual peculiarity) , as well as in a 
majority of the internal characters, it agrees well with the descriptions of Savigny 
and Hartmeyer cited above. 

But this specimen is abnormal and defective in the slight development of the 
dorsal tubercle, which is practically wanting, although its aperture, which is U- 
shaped, with the open interval obliquely forward and to the left and with one of the 
ends bent down, is clearly visible, but very small. Yet I was not able to find any 
neural duct extending from its aperture, nor any neural gland. Even the ganglion 
was only doubtfully demonstrated. The neural duct should be long in this species, 
with accessory lateral openings as well as the terminal one in the dorsal tubercle. 
The tissues of this specimen were dark coloured and somewhat opaque, but that would 
not account for the difficulty of finding the above structures if they were present in a 
normal state of development. 

5. Ascidia cannelata (Oken), 1820 

Phallusia sulcata Savigny, 1816: 162, pi. 9, fig. 2. (Name preoccupied.) 
Phallusia cannelata Oken, I sis, 1820 : 796. 
Ascidia cannelata, Hartmeyer, 1916: 400, fig. 1. 

One specimen, 32 mm. in length, growing on coral. 

6. Rhodosotna turcicum (Savigny), 1816 
Phallusia turcica Savigny, 1816: 165, pi. 10, fig. 1. 

Seven specimens, all rather small except one 45 mm. long. This, apparently the 
only species of its genus, is found in many tropical seas, and is readily recognizable 
by the two apertures being near together in a cleft of the test which can be tightly 
closed to give them protection. Said to be in most places a rather uncommon species ; 
apparently the Gulf of Aqaba is an exception, as is also the island of Curacao, West 
Indies. 

7. Cnemidocarpa hemprichi Hartmeyer, 1916 
Cnemidocarpa hemprichi Hartmeyer, 1916a: 218, figs. 6, 7. 

One specimen of very irregular external form, about 29 mm. long. Found asso- 
ciated with coral in a depth of 2 fathoms. 

8. Polycarpa mytiligera (Savigny), 1816 

Cynthia mytiligera Savigny, 1816: 158, pi. 8, fig. 2. 
Polycarpa mytiligera, Hartmeyer, 1916a: 208, figs. 1, 2. 

Two specimens, each of which contained a relatively large symbiotic macruran 
crustacean in the branchial cavity. 

9. Herdmania momus (Savigny), 1816 

Cynthia momus Savigny, 1816: 143, pi. 1, fig. 2; pi. 6, fig. 1. 
Cynthia pallida Heller, 1878: 96, pi. 3, figs. 17, 18. 

Five specimens, all of rather small size and apparently all representing the typical 
variety of this widely distributed species of warm regions. 



218 THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 

io. Microcosmus exasperatus Heller, 1878 
Microcosmus exasperatus Heller, 1878: 99, pi. 3, fig. 19. 

Three very small specimens. This is also a species of extensive distribution in 
tropical and warm-temperate waters. 

11. Halocynthia spinosa Sluiter, 1905 
Halocynthia spinosa Sluiter, 1905: 15, pi. 2, figs. 8-8d. 

Five specimens, the largest about 20 mm. in greatest diameter. 

This species, more or less red or pink in colour in life, is easily recognizable from 
its spiny exterior, the spines about the aperture on the siphons being especially long 
and conspicuously provided with sharp lateral branches. 

12. Molgula dione (Savigny), 1816 
Cynthia dione Savigny, 1816: 153, pi. 7, fig. 1. 

One specimen, about 22 mm. long, found on coral. 

Class THALIACEA 

Pelagic Tunicata 

All the Thaliacea in the collection are of one species, Salpa maxima Forskal, 1775, 
which is found in both the Atlantic and Pacific Oceans, and though reported also 
from the southern part of the Indian Ocean, has apparently not previously been 
recorded from the Red Sea. The specimens, with the possible exception of some 
immature ones as noted below, belong to the following variety of this species: 

13. Salpa maxima Forskal, 1775, var. tuberculata Metcalf, 1918 
Metcalf, 1918, Bull. U.S. Nat. Mus., No. 100, 2 (2) : 87, fig. 72. 

Described by Metcalf (who had examples of the aggregated form only, from the 
southern Philippines). The 'Manihine' collection has large adult examples of both 
aggregated and solitary forms, collected with dip nets near the surface, in some cases 
with the aid of a light. 

Five adult specimens of the aggregated form agree well with Metcalf 's description 
and figures, in having the anterior and posterior processes of the body longer than in 
the typical 5. maxima, and in having on each side of the external body surface an 
oval area of the thickened test at the base of the atrial siphon, bearing small acute 
conical spinous tubercles as described by Metcalf, the area on left side being the 
larger. 

Four adult examples of the hitherto undescribed solitary form of the variety 
tuberculata, the largest about 135 mm. in length, also differ from the solitary form 
of the typical 5. maxima in having external spinous areas, though these are small. 
There are three of these in the case of the solitary form, the most conspicuous one 
being a narrow transverse strip of thickened test extending across the rear end of the 



THE 'MANIHINE* EXPEDITION TO THE GULF OF AQABA 219 

body just below (ventral to) the base of the atrial siphon, bearing two not very regular 
rows of conical spinous tubercles similar to those in the aggregated form. The rows 
are one above the other, and extend slightly farther on the left than on the right side. 
On the dorsal surface of the body, above the intestinal 'nucleus', there is on each side 
a thickened area of test bearing a few conical tubercles, but both areas are of small 
extent, especially the one on the right side. 

The variety tuber culata appears to be a well-marked one, but the differences from 
the typical form are superficial and hardly seem to justify considering it a distinct 
species, especially since we do not yet know the extent to which intermediate forms 
may occur. 

The collection also contains a number (over 50) of young specimens of 5. maxima, 
aggregated form, measuring up to about 20 mm. in length exclusive of the anterior 
and posterior processes. Many of these, when collected, were still adhering together 
as parts of chains, but due to transportation and handling are now all separated. It 
is likely that they are all the young of the variety tuber culata, but as they fail, pro- 
bably because too young, to show the varietal characters, they have been labelled 
simply Salpa maxima. 



REFERENCES 

Forskal, P. 1775. Descriptiones animalium . . . quae in itinere orientali observavit. (Tunicata, 

pp. 112-117, 129, 130.) 
Hartmeyer, R. 1916. XJber einige Ascidien aus dem Golf von Suez. S.B. Ges. naturf. Fr. Berl. 

1915: 397-430. 14 figs. 
1916a. Neue und alte Styeliden aus der Sammlung des Berliner Museums. Mitt. zool. Mus., 

Berl., 8: 203-230, 13 figs. 
Heller, C. 1878. Beitrage zur naheren Kenntniss der Tunicaten. S.B. Akad. Wiss. Wien, 

77: 83-110, 6 pis. 
Herdman, W. A. 1882-1888. Rep. Sci. Res. Voy. H.M.S. 'Challenger', 6, 1882, Ascidiae Sim- 

plices: 1-296, 37 pis., 23 text-figs.; op. cit. 14, 1886, Ascidiae Compositae: 1-429, 49 pis., 

15 text-figs. ; op. cit. 27, 1888, Pelagic Tunicata and Appendix: 1-163, 11 pis., 28 text-figs. 
Lesueur, C. A. 1823. Descriptions of several new species of Ascidia. /. Acad. nat. Sci. Philad. 

3: 2-8, 3 pis. 
Metcalf, M. M. 1918. The Salpidae: a taxonomic study. Bull. U.S. nat. Mus., No. 100, 2 (2) : 

1-193, 150 figs., 14 pis. 
Michaelsen, W. 1919. Ascidiae Ptychobranchiae und Dictyobranchiae des Roten Meeres. 

Denkschr. Akad. Wiss. Wien, 95 (10): 1-120, 20 figs., 1 pi. 

1919a. Zur Kenntniss der Didemniden. Abh. Naturw. Hamburg, 21 (1): 1-44, 3 figs. 

1920. Ascidiae Krikobranchiae des Rothen Meeres. Denkschr. Akad. Wiss. Wien, 97 (9): 

1-38, 1 pi. 
Oken, L. 1820. [Translation into German of Savigny's work of 1816 (with the plates).] I sis 

(von Oken), 1820. 
Savigny, J. C. 1 816. Memoires sur les animaux sans vertebres, 2: 1-239, 24 pis. 
Sluiter, C. Ph. 1905. Tuniciers recueillis . . . dans la Golfe de Tadjourah. Mem. Soc. zool. Fr., 

18: 1-20, 2 pis. 
1919. "Uber einige alte und neue Ascidien aus dem Zool. Museum von Amsterdam. Bijdr. 

Dierk. 21: 1-2 1, 1 pi. 



220 THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 

APPENDIX 

Ascidian from Mukalla Bay 

Apparently Ascidia savignyi Hartmeyer, 1916 

A large specimen of the genus Ascidia from Mukalla Bay, South Arabia (A. Fraser-Brunner, 
coll. 1 7-1 2-1948) is not included in the above list of specimens as it was not from the Gulf of 
Aqaba. It is remarkable for its large size (about 160 mm. long by 35 mm. transversely) and 
greatly elongated form, due chiefly to much lengthening of the anterior half of the body, though 
the siphons (both of which arise at the anterior end) are short, and the branchial one is much 
distorted. The internal structure does not show much abnormality, though the branchial sac 
extends close to the anterior end of the body, and the dorsal tubercle (whose aperture is irregular 
S-shaped, with the upper end bent down), also the neural gland and ganglion, are close to it 
and very near to the circle of tentacles. The branchial sac has no intermediate papillae ; the 
internal longitudinal vessels are numerous (over 70 on the left and over 80 on the right side) ; 
the intestinal loop (about 37 mm. long) is far back in the body. 

It is evidently an unusually old individual ; one that has grown in a favourable position in 
respect to food-supply and protection from predatory fishes and crabs, but where surrounding 
obstructions compelled it to become unusually elongated. 

A similar specimen might be hard to find again, but I do not think it should be assumed to be 
a new and undescribed species, though such mistakes have too often been made, resulting in 
burdening literature with supposed species having no real existence. Such a specimen is hard to 
identify with certainty, but I think it is an unusually large and abnormally shaped example of 
Ascidia savignyi Hartmeyer, 191 6. (Sitzungsber . Gesell. naturf. Freunde Berlin: 1916: 404), des- 
cribed from the Sinai coast and Gulf of Suez. 

In that article Hartmeyer mentioned (p. 407) the close relationship of A. savignyi to A. 
depressiuscula Heller, 1878, described from Ceylon, and common in the Philippines, which is a 
species that also attains rather large size. I am quite ready to agree with this opinion, and think 
he was also probably correct in believing it related to the European species A . virginea Mueller, 
but I do not consider it also related to A . paratropa (Huntsman) of the American Pacific coast, 
as Hartmeyer believed. That species has intermediate papillae on the branchial sac, and belongs 
to a different section of the genus. 






IX. FISHES 

By N. B. MARSHALL, M.A. 

The collection comprises 113 species, of which 1 is new, while 4 sub-species have 
been proposed. There are n new records for the Red Sea (these being indicated by 
an asterisk preceding the name of the species). 

Collections were made from 28 December 1949 to 16 February 1950, coming from 
various localities along the Sinai shores of the gulf and from an area around the en- 
trance. These include Aqaba, Faraun Island, Graa, Mualla, Wasit, Hobeik, Dahab, 
Urn Nageila, and Abu Zabad within the gulf and Tiran Island, Sanafir Island, 
Sherm-el-Moiya, Sherm Sheikh, and Ras Muhammad Bay around the entrance. For 
the positions of these localities reference should be made to the chart in the intro- 
duction to this series of reports. 

The fishes were captured by a variety of methods: cast-net, fish-trap, hand-lines, 
trolling gear, and dip-net. In addition many were taken by bringing up pieces of 
coral and breaking them open to obtain the enclosed fishes, while a number were 
obtained from pools along the reef at low water. The method of capture is indicated 
under each species, giving certain information on the habits of the fish. For example, 
those taken by cast-net occurred singly or in shoals in shallow water close to the 
shore, while those taken by trolling spoon or live bait were nearly always caught 
along the seaward edge of reefs, where they appear to station themselves to prey on 
smaller fishes living in association with coral. Clearly those found within pieces of 
coral must live in close association with it, darting back to shelter on being disturbed 
by the diver. Perhaps no more striking way of appreciating the direct or indirect 
dependence of so many tropical fishes on coral can be obtained than through the 
many ways necessary to obtain them as specimens. 



SELACHII 
Carcharinidae 
Negaprion acutidens (Ruppell) 
1 specimen of length 660 mm. 1 taken close inshore in Ras Muhammad Bay. 

Carcharinus melanopterus (Quoy & Gaimard) 
1 specimen of 535 mm. caught by hand-line at Sanafir Island. 

Carcharinus albimarginatus (Ruppell) 
1 specimen of 870 mm. caught by hand-line at Sherm Sheikh. 

1 Except for the Selachii and the eels, lengths throughout this paper refer to the standard length. 



222 THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 

Rhinobatidae 
Rhinobatus halavi (Forskal) 

Six specimens were taken in very shallow water in Ras Muhammad Bay. One of 
these is a female of length 507 mm., while the rest are males ranging from 355 to 
520 mm. 

Dasyatidae 

Dasyatis uarnak (Forskal) 

One specimen taken by hand-line at a depth of 10 fathoms at the anchorage in 
Sanafir Island. The disk is about 1,000 mm. in length and 1,250 mm. wide. The tail, 
from which the whip-like end is missing, has a length of about 1,250 mm. 

Taeniura lymtna (Forskal) 

Three specimens were obtained by cast-net close inshore at Sanafir Island (length 
570 mm.), and at Mualla (length 445 mm.) and Um Nageila (length 564 mm.) within 
the Gulf of Aqaba. 

ISOSPONDYLI 

Clupeidae 
Sub-family dussumieriinae 

Spratelloides delicatulus (Bennett) 

Individuals of this species were taken with a dip-net and Aldis lamp at night. 
Faraun Island: 10 specimens from 21 to 50 mm. Sanafir Island: 15 specimens from 
40 to 45 mm. 

Spratelloides gracilis (Schlegel) 

Like the preceding species, this was caught by dip-net at night in the light of an 
Aldis lamp. Hundreds of specimens were taken at the anchorage at Sanafir Island, 
ranging in length from 9 to 39 mm. 

I have compared some of these specimens with material in the museum collections 
from Japan and Formosa (the type locality being along the south-east coast of 
Nagasaki). There are differences in the number of pectoral and anal fins as shown 
in the table below : 



No. of rays 

Red Sea specimens 

Japanese specimens 



Pectoral (left) 



13 14 15 
3 7 

3 6 



Anal 



11 12 13 14 

5 7 1 

262 



On the basis of the above counts it seems not unlikely that the Red Sea populations 
should be separated as a distinct sub-species ; but lacking data from areas between 
the end points of the range of this species it is considered premature to subdivide it. 



THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 223 

INIOMI 

Synodontidae 

Synodus variegatus (Lacepede) 
One specimen of 130 mm. taken in a pool at Dahab. 

APODES 

MURAENIDAE 

Echidna nebulosa (Ahl) 
Two specimens of 444 and 460 mm. taken on the reef at Abu Zabad at low tide. 

Echidna polyzona (Richardson) 

One specimen from Abu Zabad of 195 mm. and two from Sanafir Island of 115 
and 165 mm. The latter were found in a piece of madreporarian coral. 

Gymnothorax meleagris (Shaw) 

Seven specimens were obtained from the following localities: Dahab (108 mm.), 
Abu Zabad (145 and 160 mm.), Sanafir Island (in, 165, and 180 mm.), Sherm Sheikh 
(100 mm.). Except those from Abu Zabad, which were obtained on the reef at low 
tide, all were found in pieces of madreporarian coral brought up for examination. 

Gymnothorax flavimarginata (Riippell) 

One specimen of 295 mm. taken on the reef at Abu Zabad at low tide and one of 
880 mm. from Ras Muhammad Bay. 

Gymnothorax geometrica (Riippell) 

Two examples of 130 and 143 mm. taken from pieces of coral at Sherm-el-Moiya 
and Sanafir Island respectively. 

The body colour of these specimens was fawn with the pattern of dark pigment 




Fig. 1. An immature specimen of Gymnothorax geometrica 

(Riippell) from Sherm-el-Moiya in the northern Red Sea, 

showing the pattern of dark spots on the head and body. 

spots on the head looking rather like a series of lateral line pores (see Fig. 1). These 
spots extend down the mid flanks as a single line, extending just beyond the anus. 



zoo. 1. 8. 



Gg 



224 THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 

This spot pattern was also found in specimens from the collections labelled Gymno- 
thorax thyrsoidea (Richardson): 2 from Rodriguez, 1 from the Seychelles, 1 from 
Muscat. These specimens differed from those listed above in having a dotted and 
speckled body coloration, but as all these Indian Ocean examples were much larger 
it is likely to be a difference due to age. Ruppell's (1828) figure shows a mottled 
body coloration. 

There can be no doubt that G. geometrica (Riippell) and G. thyrsoidea (Richardson) 
are very closely related, the only apparent difference between them being the absence 
of the spot pattern in the latter. However, as this pattern seems quite constant in 
G. geometrica and as the pattern only appears to be found in individuals from the 
western Indian Ocean, the two species have been kept separate. Further work may 
perhaps show that what is now called geometrica is a western Indian race of a widely 
spread species. (This species will, of course, need to be called G. geometrica — this 
name having priority over G. thyrsoidea.) 

If Gymnothorax geometrica is a distinct species its distribution must include the 
western Indian Ocean as well as the Red Sea. 

Uropterygius polyspilus (Regan) 

One specimen from Sanafir Island of length 201 mm. taken in a piece of coral. 

Schultz (1943) has suggested that this species is perhaps the young of Uropterygius 
tigrinus (Lesson). Examination of the above specimen, together with the type 
specimen from Tahiti (length 183 mm.) and two specimens from Samoa (331 mm.) 
and Zanzibar (715 mm.), shows that polyspilus is distinct from tigrinus (a specimen 
of 860 mm. was examined) in the following characters: 

1. The number and size of the outer maxillary teeth: 20-28 in polyspilus, which 
are much smaller than the inner series of maxillary teeth; 12-13 in tigrinus, 
which are nearly the size of the inner series. (Bleeker, Atlas Ichthyologique, 4, 
1864: 113, counts 16 outer maxillary teeth for tigrinus. The figure on plate 
165 shows them almost equal in size to the inner series.) 

2. The proportions between trunk and tail: about equal in length in polyspilus, 
but in tigrinus the trunk is about 1-7-1-8 times longer than the tail. 

3. The proportions between the eye and the snout: in polyspilus the length of the 
snout is from 1-7 to i-8 times the diameter of the eye, whereas in tigrinus the 
snout is about 2-3 times the eye diameter. 

SYNENTOGNATHI 
Belonidae 
Strongylura crocodilus (Lesueur) 
One specimen from Sanafir Island of 465 mm. 

Hemirhamphidae 
Hemirhamphus far (Forskal) 
One specimen from Sanafir Island of 300 mm. 



THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 

EXOCOETIDAE 



225 



Danichthys rondeletii (Cuvier & Valenciennes) 

Four specimens taken off Alexandria, which were attracted on board by a light. 
Lengths 152, 164, 173, and 180 mm. Bruun (1935) has suggested that D. rondeletii 
in the Mediterranean might prove to be a dwarf race distinct from the Atlantic form. 
Examination of the above specimens, 2 others from the Mediterranean (B.M. Reg. 
No. 73.4.21.2-3) and 1 from the Atlantic (B.M. Reg. No. 71. 12.28.8) has yielded 
data which when added to those listed by Bruun and Breder (1938) provides evidence 
to support this suggestion. 

The essential differences between the two forms are in the number of pectoral rays 
and transverse scales (and very probably in the sizes attained, the Atlantic form being 
known up to 234 mm. in standard length and the Mediterranean up to 187*5 mm.). 
These differences are shown below: 



Atlantic 




Mediterranean 


specimens 


Pectoral rays 


specimens 


1 


16 


3 


11 
5 


17 
18 


9 

I 


1 


19 
Transverse scales 




A tlantic 


(between origin of 
dorsal fin and lateral 


Mediterranean 


specimens 


line) 


specimens 


2 
13 


6* 

7i 


9 

1 



It would appear from these data that the Mediterranean form can nearly always be 
separated from the Atlantic by the number of transverse scales. If more evidence, 
in particular more from the Mediterranean, shows this is so, then this species must 
be split into Atlantic and Mediterranean sub-species. 

MICROCYPRINI 

Cyprinodontidae 

Aphanius dispar (Riippell) 
One female of 32 mm. taken on the reef at Abu Zabad at low tide. 



SOLENICHTHYES 

FlSTULARIIDAE 

Fistularia villosa Klunzinger 

Four specimens from Dahab from 600 to 790 mm. Another specimen of 105 mm. 
taken with a dip-net at Sanafir Island is probably of this species. In determining this 
species I have used the revision by Duncker & Mohr (1925) and other specimens in 
the collections. 



226 THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 

The colour was noted as follows : ' a line of misty-blue spots on either side of the 
mid-dorsal line extending from the pectoral fins to the dorsal. Below this line (about 
\") a continuous misty-blue line extended from about i" in front of the pel vies to i" 
behind the end of the dorsal, thereafter continuing as a line of spots.' 

Syngnathidae 

Micrognathus brevirostris (Ruppell) 

Three specimens found in a piece of coral at Sanafir Island. Two males of 37-0 
and 47-5 mm. and one female of 44-0 mm. 

BERYCOMORPHI 

HOLOCENTRIDAE 

Holocentrum spiniferum (Forskal) 

Three specimens taken by hand-line at Sanafir Island (278 and 300 mm.) and 
Sherm Sheikh (295 mm.) at depths of about 10 fathoms. 

Holocentrum sammara (Forskal) 
One specimen from Sanafir Island (length 134 mm.). 

Holocentrum diadema Lacepede 
Two specimens of about 45 mm. from a piece of coral at Tiran Island. 

Holocentrum lacteoguttatum (Cuvier & Valenciennes) 
Two specimens of 45 and 54 mm. Taken at low tide on the reef at Abu Zabad. 

PERCOMORPHI 

(Sub-order PERCOIDEA) 

Serranidae 

(Sub-family serraninae) 

Plectropoma maculatum (Bloch) 
One specimen of 440 mm. caught by hand-line at Sanafir Island. 

Variola louti (Forskal) 

Two specimens of 385 and 287 mm. taken by hand-line at Dahab (10 fms.). This 
species could also be caught by trolling a spoon or live bait. 

Cephalopholis miniatus (Forskal) 
Two specimens of 280 and 287 mm. taken by hand-line at Dahab. 



THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 227 

Cephalopholis hemistictus (Ruppell) 

Three specimens, two from Dahab of 131 and 140 mm. and one from Hobeik of 
131 mm. 

Intensive field and laboratory work may show the two above species to be synony- 
mous. Klunzinger (1884) states that the only distinguishing feature is in the colora- 
tion, which he says is constant in hemistictus. There is, however, considerable 
variability. The usual body colour in hemistictus is brownish or dark olive-green 
with small bright blue, ocellated spots on the head and lower half of the flanks 
(mainly found on the thoracic and abdominal regions), while there is a broad yellow 
edging to the pectoral fin. The three specimens from the Gulf of Aqaba differ from 
this in the general body colour, this being a bright red as in C. miniatus. (Two other 
specimens from the Gulf of Aden and one from the Makran coast also must have 
had this coloration.) What is also interesting on all these specimens are the pale 
pelvic fins with a narrow outer black edging which is also found in C. miniatus (in 
typical C. hemistictus they have a general, dusky pigmentation). Again, the area of 
the body covered with the blue spots in hemistictus varies considerably from being 
confined to part of the abdominal region to practically extending over the lower half 
of the flanks, with a few spots appearing dorsally above the lateral line. Finally in 
five specimens labelled Epinephelus miniatus from Mombasa there are two specimens 
of 226 and 242 mm. with the normal colour pattern, while the remaining three from 
151 to 171 mm. (which agree in all characters but colour with the above two) are 
completely plain coloured. There is no trace of spots and only a faint dark edging 
to the caudal and anal fins can be seen. Presumably these were coloured a bright 
red in life. 

Although it is quite possible that these species merge with one another, they have 
been separated particularly on account of the difference in distribution, Cephalo- 
pholis hemistictus being confined to the Red Sea and western Indian Ocean, whereas 
C. miniatus occurs throughout the Indo-West-Pacific area. There is here an interest- 
ing parallel with the eels Gymnothorax geometrica and G. thyrsoidea which were 
discussed earlier in this report. 

Epinephelus summana (Forskal) 
One specimen of 440 mm. from Sanafir Island taken by hand-line. 

Epinephelus fuscoguttatus (Forskal) 

Six specimens. Four from Sanafir Island from 325 to 890 mm. caught by hand- 
line in depths from 5 to 20 fathoms. Two from Abu Zabad of 51 and 123 mm. taken 
on the reef at low tide. 

Epinephelus fasciatus (Forskal) 
Seventeen specimens taken at the following localities : Dahab, 5 from 34 to 175 mm. ; 



228 THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 

Hobeik, 3 from 190 to 220 mm. ; Abu Zabad, 7 from 46 to 76 mm. collected from pools 
on the reef at low tide ; Sanafir Island, 1 of 43 mm. ; Sherm Sheikh, 1 of 205 mm. 



(Sub-family theraponinae) 
Therapon jarbua (Forskal) 

Thirteen specimens captured by cast-net at Sherm Sheikh (12 from 79 to 116 mm.) 
and at Abu Zabad (1 of 167 mm.). 

Investigations of these specimens together with others in the collections has shown 
that there are certain regional differences in the number of dorsal spines and the 
relation between the depth of body and the standard length as shown in the follow- 
ing table : 







No. of 












dorsal spines 


depth 
X JOO 


No. of 


Size range 




Area 


11 12 


length 


specimens 


(mm.) 




Red Sea . 


13 — 


29-5-32-o 


13 


79-167 




Arabian coast . 


5 — 


32-0-35-0 


5 


80-276 




Persian Gulf . 


4 2 


33-0-38-0 


6 


58-63-5 




Coasts of India and Ceylon 


12 — 


32-1-38-6 


12 


26-113 




East African area 


2 6 


30-7-35-0 


8 


54-204 




East Indies 


1 10 


32-2-37-0 


11 


23-151 




Philippine Islands 


1 2 


36-1-37-2 


3 


90-117 




Fiji and Samoa 


— 2 


35-5-35-7 


2 


141-154 




China .... 


— 3 


31-5-32-9 


3 


34-5-96-5 




Australia 


— 2 


30-9-33-6 


2 


67-210 



Although these data are rather limited, it is clear that in the Red Sea Therapon 
jarbua has n spines in the dorsal fin (previously found by Klunzinger, 1884) and also 
tends to be slenderer in form than representatives from the Indo-Pacific areas. 
Furthermore, the proportion of the Indian Ocean specimens having n as against 12 
spines is 23 : 8, whereas in the Pacific Ocean this is 2 : 19. Of specimens from the 
Pacific, those from the Philippines, Fiji, and Samoa have the deepest body form. 

It is thus quite evident that the populations of Therapon jarbua are by no means 
uniform in character. Whether, for example, the Red Sea population can be con- 
sidered to be part of a sub-species found mainly in the north-west Indian Ocean 
(having 11 dorsal spines), which intergrades over a wide area with a typical Pacific 
sub-species (having 12 dorsal spines), can hardly be decided on the present data. 
It is, however, a problem worth much further investigation. 

During this work it became necessary to decide whether Therapon servus (Bloch) 
is distinct from T. jarbua (Forskal). Weber & de Beaufort (1931) have synonymized 
them but refer to the work of Jordan & Thompson (1912), who decided that they 
were good species, particularly separated by the longitudinal scale counts just above 
the lateral line. The present work confirms Jordan & Thompson's conclusions and 
shows that in general Therapon servus has relatively smaller scales than T. jarbua, 
as shown in the following table. 



THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 



229 



Scale count 


Therapon jarbua 


No. specimens 
seen 


Therapon servus 


No. specimens 
seen 


1. Longitudinal series above 
the lateral line 

2. Transverse scales 

3. Rows of scales on preo- 
perculum 


77-89 
(12) 14-17 
25-30 

8-1 1 


57 
55 

57 


92-105 
17-21 
3o-35 

11-13 


12 
12 

12 



Serranidae 

Sub-family grammistinae 

Grammistes sexlineatus (Thunberg) 
Three specimens from 70 to 82 mm. taken at low tide on the reef at Abu Zabad. 

Sub-family pseudochromidinae 

Pseudochromis olivaceus Ruppell 

All the examples of this species were taken from pieces of coral brought up by a 
diver. Within the Gulf of Aqaba collections were made at Graa (2 specimens of 
26 and 45 mm.), Mualla (4 specimens of 23-47 mm.), and at Dahab (4 specimens from 
37 to 59 mm.). There are also 34 from 26 to 70 mm. taken at Sanafir Island and 
8 from 29 to 54 mm. taken at Sherm-el-Moiya. 

Comparison has been made between the Gulf of Aqaba individuals and some of 
those taken outside the entrance in the Red Sea. There does appear to be some 
difference in the number of pectoral rays, which are tabulated below : 



Pectoral rays 


17 


18 


19 


Gulf of Aqaba 
Sanafir Island 


3 


6 
10 


1 
2 



This species is confined to the Red Sea. 

Plesiopidae 
Plesiops nigricans (Ruppell) 
Twenty-three specimens from ^ to 63-5 mm. collected at Abu Zabad at low tide. 

Ciieilodipteridae 

Apogon endekataenia Bleeker 

Nine specimens from Abu Zabad from 53 to 61 mm., collected on the reef at low 
tide. 

These specimens agree in structure with two specimens in the collections (labelled 
as types) which were obtained from Bleeker (B.M. Reg. No. 1880. 4. 21. 59-60). The 
latter have nearly lost all trace of colour but still retain the remains of the spot on 



230 THE 'MANIHINE* EXPEDITION TO THE GULF OF AQABA 

the base of the caudal fin which Weber & de Beaufort (1929) list as one of the charac- 
ters separating A. endekataenia from A. novemfasciatus C.V. Comparison of these 
specimens with those of novemfasciatus shows the two to be very distinct in tooth 
character. In the latter the teeth are relatively large, there being 4 rows in the 
upper and lower jaws while in endekataenia there are from 6 to 9 somewhat irregular 
rows of smaller teeth. Comparison of specimens of equal size shows that the teeth of 
novemfasciatus are about twice the size of those of endekataenia. 

Examination of the museum collections has not revealed any examples of A. 
novemfasciatus from the Red Sea or Indian Ocean. Klunzinger (1884) notes that his 
specimens (which he names A . fasciatus White) show clearly the black spot on the 
base of the tail. Smith (1949), however, records it as quite common north of Zulu- 
land. 

Cheilodipterus quinquelineatus Cuvier & Valenciennes 

Three specimens from 31 to 38 mm. taken at Abu Zabad. 

Latilidae 

*Malacanthus hoedtii Bleeker 
One specimen from Sherm Sheikh of 207 mm. 

Carangidae 

Caranx fulvoguttatus (Forskal) 
One specimen from Sanafir Island of 170 mm. 

Caranx sexfasciatus Quoy & Gaimard 
Two specimens of 544 and 800 mm. caught by trolling a spinner at Sanafir Island. 

Lutianidae 

Lutianus bohar (Forskal) 

Two specimens caught by hand-line at Sanafir Island (length 310 mm. ; depth 
20 fms.) and at Sherm Sheikh (length 357 mm. ; depth 6 fms.). 

Lutianus argentimaculatus (Forskal) 

One specimen of 345 mm. caught by hand-line at a depth of 20 fms. at Sanafir 
Island. 

Lutianus fulviflamma (Forskal) 

Three specimens caught by hand-line (two from Sanafir Island of 222 and 232 mm. 
taken at 20 and 8 fms. respectively ; one from Sherm Sheikh of 209 mm. from 6 fms.) 

Lutianus kasmira (Forskal) 

Six specimens taken on hand-lines Four from Sherm Sheikh at 147-182 mm. and 
two from Hobeik at 209 and 211 mm. 



THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 231 

Aphareus rutilans Cuvier & Valenciennes 

One specimen of 765 mm. obtained from the cold store at Aqaba. 

This is one of the finest food fishes taken in the Gulf of Aqaba and is known to the 
Arab fishermen as Faris. It is caught by hand-line mainly at depths of about 100 
metres. 

Mullidae 

Parupeneus tnacronema (Lacepede) 

Five specimens. Three obtained by cast-net (two at Dahab of 145 and 149 mm. 
and one at Sanafir Island of 96 mm.). The other two of 83 and 84 mm. were caught 
in a fish-trap at Aqaba at a depth of 10 fathoms. 

Lethrinidae 

Lethrinus nebulosus (Forskal) 

Two specimens of 320 and 450 mm. caught by hand-line at Sanafir Island (5 fms.) 
and Dahab (15 fms.) respectively. 

Lethrinus tnahsena (Forskal) 

Four specimens taken at Dahab (two of 290 and 310 mm. at 12 fms.) and Sanafir 
Island (two of 225 and 257 mm. at 5 fms.). 

Lethrinus microdon Cuvier & Valenciennes 

Five specimens, of which three are from Aqaba (86-97 mm.) taken in a fish-trap 
at a depth of 10 fathoms. The other two were caught by hand-line at Dahab (length 
317 mm. ; depth of water 7 fms.) and Sanafir Island (length 360 mm. ; depth 5 fms.). 

Lethrinus mahsenoides ([Ehrenberg] Cuvier & Valenciennes) 

Twelve specimens. Seven from Aqaba taken in a fish-trap set at 10 fathoms. Three 
from Dahab of 176, 183, and 184 mm. caught by hand-line at a depth of 10 fathoms. 
One from Hobeik of 200 mm. from a depth of 10 fathoms, and one from Sherm 
Sheikh of 162 mm. from 6 fathoms. 

Weber & de Beaufort (1936) have remarked that L. mahsenoides from the Red Sea 
is hardly separable from L. ornatus C.V. (= L. insulindicus Bleeker). I have com- 
pared the above specimens and one of Klunzinger's from the Red Sea (labelled 
mahsenoides) with those labelled 'mahsenoides* and insulindicus taken outside the 
Red Sea. I could find no significant differences. 

Gymnocranius griseus (Schlegel) 

One specimen from Hobeik of 300 mm. taken by hand-line at a depth of 10 fathoms. 

The above specimen has been compared with two from Mauritius (B.M. Reg. Nos. 
1932.8.8.22 and 1934.2.22.25) and one from the Loyalty Islands (77.7.24.2), but there 
appear to be no differences. Specimens from nearer the type locality (SW. coasts of 

zoo . 1. 8. Hh 



232 THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 

Japan) differ from the Red Sea and Indian Ocean examples in being deeper bodied 
(these were from Hong Kong, B.M. Reg. No. 1939. 1. 17. 38 and the Inland Sea of 
Japan, B.M. Reg. No. 1907. 12. 23. 230-1). The depth in these is about half the 
standard length as against T 5 j to ^ in the Red Sea and Mauritius specimens. There is, 
also, a difference in coloration, for the Red Sea and Indian Ocean specimens have 
the wavy blue lines across the head, a coloration which never seems to be present in 
Pacific Ocean fishes of this species. Fowler (1933) has even made this difference the 
basis for two sub-genera. 



Sparidae 
Sparus bifasciatus (Forskal) 

Two specimens from Sanafir Island (92-5 mm.) and from Um Nageila (154-0 mm.). 

On comparing these specimens with others in the museum collections it became 
quite evident that there are two definite colour varieties. The first, which is found 
in the Red Sea, the Gulf of Aden, along the South Arabian coast (Muscat), and in 
the Persian Gulf, has plain hyaline or yellow dorsal and caudal fins. The other from 
the Makran coast of Baluchistan, the north-western Indian coast, and the East 
African area (specimens from Kosi Bay, Zululand, and Rodriguez) always has a 
black edging to the dorsal fin and sometimes a black edging in the fork of the caudal. 
Reference to the literature on this species confirms this difference in pigmentation 
and the geographical range of each type. 

In body proportions and height and lengths of the fins there are no significant 
differences between these two forms. In fin ray and scale counts there are also no 
differences, except that there appears to be a definite tendency for the East African 
examples to have 13 rays in the soft dorsal rather than 12. Smith (1938) also gives 
13 as the number of dorsal rays in a specimen from Natal. Counting the latter, five 
out of six East African examples have 13 rays, whereas from the rest of the area of 
distribution only one out of eighteen had this number; the rest had 12. 

It is not the intention to do more at this stage than draw attention to this differ- 
entiation within the populations of this bream. More data on the Baluchistan and 
north-west India populations would be of interest, for at present it appears that, 
although they have the same colour pattern as the East African, they tend to have 
12 dorsal rays rather than 13 (5 out of 6 examined). Yet one specimen from this 
area did have 13 rays. It is of interest to note that in all instances this number was 
associated with a black-edged dorsal fin. 

Sparus haffara Forskal 
Two specimens of 165 and 172 mm. taken by cast-net at Sanafir Island. 

Argyrops spinifer (Forskal) 
One specimen of 357 mm. caught by hand-line at Dahab at a depth of 10 fathoms. 



THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 233 

Diplodus noct ([Ehrenberg] Cuvier & Valenciennes) 

Ten specimens taken by cast-net at the following localities: Dahab (2 of 66-o 
and 136 mm.), Abu Zabad (2 of 140 and 146 mm.), Sanafir Island (6 from 76-0 to 
88-o mm.). 

The distribution of this species is given as the Red Sea, the Arabian and Indian 
coasts, and Madagascar (Fowler, 1933). 

Close comparison of the above material with specimens labelled Diplodus nod from 
Karachi (1) and from the Persian Gulf (n) (from Bushire) has shown them to be 
quite different. The latter are actually Diplodus sargus (Linnaeus). They are, in 
fact, the same fish as another series labelled Diplodus capensis from Muscat, Arabia, 
the latter being a synonym of D. sargus. 

The characters showing the differences between Diplodus nod from the Red Sea 
and D. sargus from the Persian Gulf and north-west Indian coast are listed below. 
The measurements and counts on D. nod were made on the 10 specimens listed above 
and 1 other of length 212 mm. from Klunzinger's collection, while those on D. sargus 
were obtained from the n specimens from the Persian Gulf (ranging in length from 
62-0 to 130-0 mm.), 1 from Karachi (of 109 mm.), and 4 from Muscat (from 140 to 
213 mm.). 

Diplodus nod (Ehrenberg) (C.V.). The greatest depth of the body is from 39-0 
to 42-1 per cent, of the standard length. Dorsal XII. 12-14 (5 specimens with 13 
rays, 4 with 14, 1 with 12). Anal III. 12-13 (5 with 12 rays, 6 with 13). Scale count 
above and below lateral line 6-7/ 15-16. Number of gill rakers on 1st arch 6-7+1 + 
12-13. 

Diplodus sargus (L.). The greatest depth of the body is from 45 to 50 per cent, of 
the standard length. Dorsal XII. 13-15 (2 specimens with 13 rays, 10 with 14, and 
2 with 15). Anal 12-14 (2 specimens with 12 rays, 9 with 13, and 4 with 14). Scale 
count above and below the lateral line 8-9/15-18. Number of gill rakers on 1st arch 
6+1+9-10. 

Reference to the literature suggests, in conjunction with the above data, that 
D. nod is confined to the Red Sea. Day (1875) records this species from the Red 
Sea and Sind (NW. India). His synopsis (p. 133) fits very well with the characters 
listed above for D. nod and his figure (pi. 32, fig. 5) is almost certainly drawn from a 
specimen of nod. Unfortunately he does not state the locality of this specimen, 
but does mention that this fish is common at Suez. His specimens from NW. India 
may well have been D. sargus. 

Sargus kotschyi Steindachner from the Arabian Gulf, Madagascar, which is 
synonymized with Diplodus nod by Fowler (1933), is probably a synonym of D. 
sargus. In particular the number of scale rows (8) above the lateral line is a good 
indication. 

In the course of this work specimens of Diplodus sargus from the Mediterranean 
were compared with those from Muscat and the Persian Gulf and good agreement 
found between them. The only difference found was in the number of scale rows 
above the lateral line, which in the Mediterranean examples was 7 to 8 compared to 
8 to 9 in those from the Arabian area. It is hoped at a later date to investigate the 
degree of differentiation within this species. 



234 THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 

Crenidens crenidens (Forskal) 

Twelve specimens taken by cast-net at the following localities : Dahab (8 specimens 
from 68-5 to 95 mm.), Sanafir Island (4 specimens from 107 to 120 mm.). 

Comparison of these specimens with others from Aden (6 collected by Mr. A. Fraser- 
Brunner) and Karachi (13) has shown that this species can be divided into two sub- 
species. 

The first is typified by specimens from the Red Sea. The diagnosis which follows 
is based on the 12 specimens listed above, 1 of 123 mm. from the Red Sea (Ruppell's 
collection), Ismailia (Suez Canal) (1 of 152 mm.), Korbrat, Suez (1 of 109 mm.), and 
the Gulf of Suez (1 of 95 mm.). The latter three specimens were collected by the 
Cambridge Expedition to the Suez Canal, 1924. 

Crenidens crenidens crenidens (Forskal) 

Depth of body 33*3-38-9 per cent., depth of caudal peduncle 9-9-10-9 per cent., 
height of third dorsal ray 9- 6-1 1 -i per cent., and length of pelvic fin 18-1-21-2 percent, 
of the standard length. Rows of scales above lateral line (from origin of dorsal) 5-6 
(7 specimens with 5 rows and 9 with 6 rows). Rows of scales below lateral line 11-12 
(2 specimens with n rows and 14 with 12 rows). Red Sea. 

Synonymy. Presumably all references to Crenidens crenidens (Forskal) or Creni- 
dens forskdlii C.V. from Red Sea localities must come under this sub-species. 

Sparus crenidens Forskal, 1775, Descript. Animal.: 15 (type locality Red Sea: Djidda or Suez). 

Crenidens crenidens, Norman, 1927, Trans, zool. Soc. Lond. 22: 380. 

Crenidens forskdlii, Cuvier & Valenciennes, 1830, Hist. Nat. Poiss. 6: 378, pi. 162 quater (type 
locality: Red Sea). Gunther, 1859, (partim) Cat. Fish. Brit. Mus. 1: 424. Klunzinger, 1870, 
Verh. zool. hot. Ges. Wien, 20: 748. Day, 1875, (partim) Fishes of India, 1: 133. 

Crenidens for skaelii, Day 1889 (partim) Fauna British India 2: 35. 

The second sub-species is typified by a series of 13 specimens from Karachi ranging 
in length from 52-5 to 164 mm. The following diagnosis is based on these individuals. 

Crenidens crenidens indicus Day 

Depth of body 43-3-49-1 per cent., depth of caudal peduncle 11-4-12-8 per cent., 
height of third dorsal ray 11-6-13-9 per cent., and length of pelvic fin 20-7-25-2 per 
cent, of standard length. Rows of scales above the lateral line 6-7 (3 specimens 
with 6 rows and 10 with 7 rows). Rows of scales below the lateral line 12-15 (3 
specimens with 12 rows, 2 with 13 rows, 7 with 14 rows, and 1 with 15 rows). Karachi. 

Synonymy. 

Crenidens indicus, Day, 1873. The sea-fishes of India and Burma from Report on the sea fish and 
fisheries, p. clxxxvi, No. 184. Day, 1875, Fishes of India, pt. 1: 132, pi. 32, fig. 4. Day, 
1889, Fauna of British India, 2: 34, fig. 13. Steindachner, 1907, Denkschr. Akad. Wiss. Wien. 
71 (1): 136. Blegvad, 1944, Danish Sci. Inv. Iran, 3: 143, fig. 80, pi. viii, fig. 3. 

Crenidens macr acanthus, Gunther, 1874. Ann. Mag. nat. Hist. (4) 14: 368 (type locality: Madras). 

Of particular interest are the six specimens from Aden mentioned above which 
range from 127 to 167 mm. in length. These have the following proportions and 



THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 235 

counts: Depth of body, 397-44-0 per cent., depth of caudal peduncle 11-1-12-0 
per cent., height of third dorsal ray 9-0-11-0 per cent., and length of pelvic fin 
20-3-21-5 per cent, of the standard length. Scale rows 6-12 (13 in one specimen). 

It will be seen that these individuals resemble Crenidens crenidens crenidens in the 
relative height of the third dorsal ray and the scale counts, but in depth of body, 
caudal peduncle, and length of pelvic fin they are more like C. c. indicus. It was the 
examination of these intermediate specimens which partly suggested the differentia- 
tion of C. crenidens into Red Sea and Arabian Sea sub-species. 

As the diagnosis shows, the latter sub-species indicus is quite distinct along the 
north-west coast of India and seemingly in the Iranian Gulf, to judge from pi. viii, 
fig. 3, in Blegvad's report (1944, loc. cit.). More specimens from the south Arabian 
coasts are clearly required. 

There is also little comprehensive data from the East African area. Two specimens 
from Mombasa and Port Natal of 118 and 186 mm. respectively closely correspond 
with C. c. indicus in body proportions, but like C. c. crenidens have 6 and 12 rows of 
scales above and below the lateral line. On the other hand, the accurate figures of 
Smith (1938, fig. 21, and 1949, pi. 44, fig. 732), together with the descriptions, give 
much more the impression of C. c. crenidens. It is thus evident that many more 
specimens from this area must be studied before the C. crenidens complex can be 
more fully appreciated. 

Pempheridae 

Pempheris sp. (probably P. moluca C.V.) 

Twenty-five juvenile specimens from 17 to 23 mm. caught by dip-net close inshore 
at Faraun Island. 

Chaetodontidae 

Chaetodon fasciatus Forskal 

One specimen of 88 mm. caught by cast-net around coral at Sanafir Island. 
This species is confined to the Red Sea. 

Anisochaetodon auriga (Forskal) 

Three specimens of 43, 48, and 51 mm. taken by cast-net at Sanafir Island. 
None of these examples have the elongated fifth or sixth dorsal ray. The two smaller 
specimens have a round black spot towards the ' apex ' of the dorsal fin. 

Platax orbicularis (Forskal) 

Nine specimens from 64 to 84 mm. taken by cast-net at Sanafir Island. 

The above individuals together with two more from the Red Sea have been com- 
pared with examples from the Indian and Pacific Oceans (Ceylon (2), Seychelles (1), 
Mombasa (1), Singapore (1), Borneo (2), Philippines (2), Manado (3), and the coast 
of Savaii (1)). 



236 THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 

There appear to be no differences except in the number of pectoral rays (counted 
in the left fin). 



Pectoral rays 


16 


17 


18 


19 


Red Sea 
Indo-Pacific . 


3 

i 


7 
4 


1 
7 


1 



POMACENTRIDAE 

Amphiprion bicinctus Riippell 
One specimen of 52 mm. taken by dip-net among coral at Dahab. 

Abudefduf biocellatus (Quoy & Gaimard) 

Eighteen specimens from 34 to 62 mm. taken on the reef at Abu Zabad at low tide. 
Three of the above have the typical biocellatus colour pattern : the rest have only 
the posterior ocellus at the base of the last few dorsal spines. 

Abudefduf sordidus (Forskal) 
Three specimens of Sy, 119, and 123 mm. caught by cast-net around rocks. 

Chromis coeruleus (Cuvier & Valenciennes) 

Forty-eight specimens, all taken from pieces of coral obtained by a diver at the 
following localities : Sanafir Island, 36 from 22 to 44 mm. ; Sherm Sheikh, 9 from 27 
to 34 mm. ; Dahab, 3 from 18 to 36 mm. 

Dascyllus aruanus (Linnaeus) 

Forty-four specimens obtained from pieces of coral at the following localities: 
Sanafir Island, 38 from 17 to 50 mm. ; Graa, 3 from 28 to 32 mm. ; Dahab, 3 from 

40 to 46 mm. 

Dascyllus marginatus (Riippell) 

Five specimens from 20-5 to 36-0 mm. obtained from a piece of coral at Dahab 
(depth 25 fms.). 

Comparison of these specimens and others from the Red Sea with those from 
localities in the Indian and Pacific Oceans has shown that a separate sub-species may 
occur in the Red Sea. A description of the diagnostic features follows below, based 
on the five specimens listed above, 1 from the northern Red Sea, taken off the Gulf 
of Aqaba (length 38-0 mm.), B.M. Reg. No. 1938. 1.24.3; 2 from the Red Sea (of 39 
and 42 mm.), B.M. Reg. No. 1935. 9. 1.5; 3 from the Kamaran Islands (from 32 to 

41 mm.), B.M. Reg. No. 1937.4.26. 8. 10; and 18 from Massaua (from 24 to 44 mm.), 
B.M. Reg. No. 71.4.13.40. 



THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 237 

Dascy litis marginatus marginatus (Riippell) 
(Fig. 2a) 

Length of longest dorsal ray (usually the fifth) from 21-9 to 287 per cent, of the 
standard length (mean 23-8 per cent. ; length of longest anal ray (usually the fourth) 
from 22-5 to 28-0 per cent, of the standard length (mean 25-3 per cent.). Rays in 
left pectoral fin (17) 18-19 ( 20 ) ( 2 specimens with 17 rays, 5 with 18 rays, 21 with 




Fig. ia. Dascyllus marginatus marginatus. Locality: Dahab, Gulf 

of Aqaba. 

Fig. ib. Dascyllus marginatus reticulatus. Locality: Philippine 

Islands. 

Fig. 2c. A specimen of D. marginatus from Aden — intermediate in 
certain respects between the two above sub-species. 

19 rays, and 1 with 20 rays). General body colour pallid to brownish (in spirits) with 
the anterior half to two-thirds of the trunk usually tending to be darker in colour 
than the rest of the body. Upper third to a half of spinous dorsal black ; this edging 
continuing along the soft dorsal as a rather thinner band as far as the tips of the 
longest dorsal rays. Anal fin with membranes between the spinous and first 5 or 6 
soft rays coloured black, contrasting sharply with the posterior half of the fin where 
the fin membranes are translucent. 
Distribution. Red Sea. 

Synonymy. 

Pomacentrus marginatus Riippell, 1828. Atlas Reise nordl. Afrika. Fische des Rothen Meers.: 38. 

pi. 8, fig. 2 (type locality: Massaua, Red Sea). 
Dascyllus marginatus Cuvier & Valenciennes, 1830. Hist. Nat. Poiss. 5: 439, pi. 133. Giinther, 



238 THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 

1862, Cat. Fish. Brit. Mus. 4: 14. Klunzinger, 1871, Verh. zool. hot. Ges. Wien 21: 520. Koss- 
man & Rauber, 1877. Zool. Ergebn. K. Acad. Wiss. Berlin, 1: 23. Borsieri, 1904, Ann. Mus. 
Civ. Genova (3) 1 (41): 214. Bamber, 1915, /. linn. Soc. Lond. 31: 481. 

The other material studied was as follows: 

Specimens from the Gulf of Aden collected by Mr. A. Fraser-Brunner, 3 specimens 
from Alayu, British Somaliland (from 30-5 to 34-0 mm.) ; 5 specimens from Berbera, 
British Somaliland (from 27-5 to 35-5 mm.) ; 1 specimen from Perim (of 35 mm.) ; 
1 from Aden (of 33-5 mm.) and 1 from Burum near Mukalla, Indian Ocean ; 2 from 
Zanzibar, B.M. Reg. No. 64.11. 15. 100 and 65.3.18.35 (of 48-0 and 52-5 mm.) ; Pacific 
Ocean 1 ; 3 from Rotuma, B.M. Reg. No. 97.8.23. 141-143 (from 26-0 to 63-5 mm.) 1 ; 
1 from Borneo, B.M. Reg. No. 58.4.21.363 (of 42 mm.) 1 ; 1 from Ponape, B.M. Reg. 
No. 76.5.19.7 (of 31-0 mm.), and 8 from Duquamete, Or Negros, Philippine Islands, 
B.M. Reg. No. 1933.3. 1 1.440-7 (from 25-0 to 58-0 mm.). The type specimen of 
Dascyllus nigripinnis Regan was also examined (B.M. Reg. No. 1908.3.23.98). 

Dascyllus marginatus reticulatus (Richardson) 
(Fig. 2b) 

Length of longest dorsal ray from 19-8 to 23-2 per cent, of the standard length 
(mean 21-1 per cent.). Length of longest anal ray from 19-1 to 24-5 per cent, of the 
standard length (mean 21-3 per cent.). Rays in left pectoral fin (19) 20-21 (1 specimen 
with 19 rays, 7 with 20 rays, and 8 with 21 rays). 

General body colour brown to dark brown (in spirits), the darker edging of the 
scales often showing up as a reticulated pattern over the body. Spinous dorsal fin 
dark brown, this pigmentation not extending to the longest rays of the soft dorsal. 
Anal fin uniformly dark brown, although sometimes the distal half of the fin may 
appear lighter. 

Distribution. Indo-West Pacific area (excluding the Red Sea). 
Synonymy. This is not complete, but lists all the names which have been proposed 

for the Indo-Pacific individuals of this sub-species. 

Heliases reticulatus, Richardson, 1845 (1846), Rep. Brit. Ass. Adv. Sci. Ichth. China &> Japan: 

254 (type locality: China Seas). 
Tetradrachmum reticulatum, Bleeker, (1872), Ned. Tijdschr. Dierk. 2: 145. 
Dascyllus xanthosoma, Bleeker, 1851, Natuurk. Tijdschr. Ned.-Ind. 2: 247. 
Dascyllus marginatus, Play fair & Gunther, 1866, Fishes of Zanzibar, 277: 81. 
Pomacentrus unifasciatus, Kner, 1868, S.B. Akad. Wiss. Wien, 58 (1): 31, 348, pi. 8, fig. 24. 
Dascyllus nigripinnis, Regan, 1907, Trans, linn. Soc. Lond. Zool. (2) 12: 228, pi. 24, fig. 5. Type 

locality: Maldives. 
Dascyllus trimaculatus (non Riippell), Fowler, 191 8, Copeia, 58: 64. 

Finally the specimens from the Gulf of Aden were found to have the following 
characteristics : 

Length of longest dorsal ray 21-4-24-6 per cent, of standard length (mean 22-7 
per cent.). Length of longest anal ray 20-6-25-9 per cent, of standard length (mean 
22-3 per cent.). Rays in left pectoral 17-19 (1 with 17, 2 with 18, and 8 with 19 rays). 
Colour in spirits dark purple-brown to brown with the caudal peduncle and the region 

1 These are labelled Dascyllus xanthosoma. 



THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 239 

over the dorsal half of the body and below the dorsal fin lighter in colour. Distal half 
to two-thirds of spinous dorsal black, this continuing as a thin edging to the soft 
dorsal as far as the tips of the largest rays. Anal, except for a lighter posterior 
edging, brownish black (see fig. 2c). 

It will be seen that these specimens are in certain respects intermediate between 
the two sub-species described above. In colour they are much like D. m. reticulatus, 
although that of the dorsal fin is more like D. m. marginatus. 

In number of pectoral rays they are clearly closest to marginatus, but are perhaps 
intermediate in the height of the longest dorsal and anal rays. The existence of inter- 
mediate forms in the Gulf of Aden suggests that this is an area where the two sub- 
species meet and interbreed. Much more material is required, however, from both the 
southern end of the Red Sea and the Gulf of Aden to establish the interrelations of 
the sub-species. 

Labridae 

Labroides dimidiatus (Cuvier & Valenciennes) 

One specimen of 27 mm. caught by hand-net among coral at Mualla. 

Thalassoma giintheri (Bleeker) 

Four specimens caught by hand-line at the following localities: Sanafir Island, 
2 of 105 and 107 mm. ; Tiran Island, 1 of 102 mm. ; and Sherm Sheikh, 1 of 154 mm. 

Thalassoma lunare (Linnaeus) 
Two specimens of 115 and 151 mm. caught by hand-line at Ras Muhammad Bay. 

Stethojulis axillaris (Quoy & Gaimard) 
Two specimens of 53 and 66 mm. taken at Abu Zabad at low tide on the reef. 

Stethojulis albovittata (Bonnaterre) 
One specimen of 79 mm. taken at Abu Zabad at low tide on the reef. 

*Halichoeres margaritaceus (Cuvier & Valenciennes) 
Three specimens of 37, 43, and 51 mm. taken at low tide on the reef at Abu Zabad. 

Cheilinus mentalis Riippell 

Eight specimens from 55 to 8y mm. taken at Aqaba in a fish trap set at 10 fathoms. 
De Beaufort (1940) has correctly synonymized Cheilinus orientalis Gunther with 
this species. There are no differences between the above specimens and the type 
specimen (B.M. Reg. No. 1864.5. 15. 8). 

Pseudocheilinus hexataenia (Bleeker) 

Four specimens all taken from pieces of coral. Three from Sanafir Island of 19, 
23, and 29 mm. and one from Sherm Sheikh of 22-5 mm. 
zoo. 1.8. ii 



2 4 o THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 

SCARIDAE 

Leptoscarus vaigiensis (Quoy & Gaimard) 
One specimen of ioo mm. collected on the reef at Abu Zabad at low tide. 

Sub-order ACANTHUROIDEA 

Acanthurus nigrofuscus (Forskal) 

Three specimens collected at Mualla by cast-net (2 of 57 and 86 mm.) and at Abu 
Zabad at low tide (1 of 55 mm.). 

Sub-order TEUTHIDOIDEA 

Teuthis rivulatus (Forskal) 

Five specimens taken by cast-net at Um Nageila (3 of 217, 220, and 235 mm.) and 
Sanafir Island (2 of 130 and 172 mm.). 

Sub-order SCOMBROIDEA 

Thynnus (Neothunnus) albacora (Lowe) 

One specimen of 1,070 mm. obtained from Arab fishermen who were catching this 
fish and the one following at a depth of about 100 metres, a few miles south of Aqaba. 

Euthynnus (Katsuwonus) pelamis (Linnaeus) 
One specimen of 670 mm. obtained a few miles south of Aqaba from local fishermen. 

Scomberomorus commerson (Lacepede) 
One specimen of 860 mm. from Sanafir Island, caught by trolling spoon-bait. 

Sub-order GOBIOIDEA 
Eleotridae 

*Eviota gymnocephalus M. Weber 

Fourteen specimens, all obtained from pieces of coral brought up by a diver 
(5 from Sanafir Island from io-o to 16-0 mm. ; 4 from Sherm Sheikh from 8-0 to 
14-0 mm. ; 1 from Sherm-el-Moiya of 15-0 mm. ; 2 from Graa of 9-5 and io-o mm.; 
and 2 from Dahab of 13-0 and 15-5 mm.). 

*Eviota distigma Jordan and Seale 

Four specimens obtained from pieces of coral at Sherm Sheikh (3 from 13-0 to 
17-0 mm.) and Graa (1 of 14-0 mm.). 



THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 241 

Hetereleotris vulgar e (Klunzinger) 

Eighteen specimens collected from pieces of coral at the following localities: 
Sanafir Island (10 from 18-0 to 26-0 mm.), Tiran Island (1 of 17-0 mm.), Mualla (3 from 
18*0 to 23-0 mm.), Dahab (4 from 19-0 to 25-0 mm.). 

This species has only been recorded from the Red Sea. 

Klunzinger (1870) remarks that the body of this fish appears to be without scales. 
I was also unable to find any trace of scales. 

Gobiidae 
Bathygobius fuscus (Ruppell) 

Three specimens collected at Dahab (1 of 34-0 mm.), Mualla (1 of 50-0 mm.), and 
Abu Zabad (1 of 47-0 mm.). All were taken close inshore where they were found 
under stones and rocks. 

Acentrogobius ornatus (Ruppell) 
Two specimens of 38-0 and 58-0 mm. taken under stones at Abu Zabad at low tide. 

Gobiodon quinquestrigatus (Cuvier & Valenciennes) 

Forty-four specimens, all obtained from pieces of coral at the following localities : 
Dahab (7 from 16-5 to 38-0 mm.), Sanafir Island (24 from 12*5 to 38*0 mm.), Tiran 
Island (10 from 16-0 to 37-0 mm.), and Sherm Sheikh (3 from 22-0 to 30-0 mm.). 

* Gobiodon erythrospilus Bleeker 

Three specimens obtained from coral at Dahab (2 of 34-0 and 37-0 mm.) and 
Tiran Island (1 of 32-0 mm.). 

Gobiodon citrinus (Ruppell) 

Six specimens obtained from pieces of coral at Sanafir Island (4 from 27-5 to 
32-0 mm.) and Sherm Sheikh (2 of 26-0 mm.). 

Paragobiodon echinocephalus (Ruppell) 

Three specimens from 20*0 to 23-0 mm. taken from a piece of coral at Sanafir 
Island. 

Sub-order BLENNIOIDEA 

Blenniidae 

Enchelyurus kraussii (Klunzinger) 

One specimen of 30 mm. taken from a piece of coral at Graa. This species has only 
been recorded from the Red Sea. 

Cirripectus variolosus (Cuvier & Valenciennes) 
One specimen of 31-0 mm. collected on the reef at Abu Zabad. 



242 THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 

Istiblennius edentulus (Bloch & Schneider) 

Twenty-five specimens collected under stones and rocks at Abu Zabad (15 from 
46-0 to 84-0 mm.) and Dahab (10 from 25-0 to 57-0 mm.). 

Istiblennius fasciatus (Bloch) 
Two specimens from Abu Zabad (1 of 47-0 mm.) and Sanafir Island (1 of 48-0 mm.). 

CONGROGADIDAE 

Haliophis guttatus (Forskal) 

Four specimens obtained from pieces of coral at Sanafir Island (3 of 50*0, 67-0, 
and 8i-o mm.) and Sherm Sheikh (1 of 6o-o mm.). 
This species appears to be restricted to the Red Sea. 

Clinidae 

*Helcogramma trigloides (Bleeker) 
One specimen of 27-0 mm. found in a piece of coral at Mualla. 

Sub-order MUGILOIDEA 

Sphyraenidae 

Sphyraena jello Cuvier & Valenciennes 
One specimen of 530 mm. taken by trolling spoon-bait at Sanafir Island. 

Sphyraena picuda Bloch. Schneider 
One specimen of 760 mm. taken by spoon-bait at Sanafir Island. 

Mugilidae 

Oedalechilus labiosus (Cuvier & Valenciennes) 

Eight specimens taken by cast-net at Mualla (2 of 101 and 104 mm.) and Sherm 
Sheikh (6 from 96 to 127 mm.). 

Liza seheli (Forskal) 
One specimen of 300 mm. taken by cast-net at Dahab. 

Liza crenilabis (Forskal) 

One specimen of 69-5 mm. from Dahab and eight specimens from 10 1 to 164 mm. 
from Sanafir Island taken by cast-net. 

Atherinidae 

Hypoatherina gobio (Klunzinger) 
Twenty-nine specimens caught by dip-net and a light at night-time at the following 



THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 243 

localities: Dahab (7 from 26-0 to 82-0 mm.), Sanafir Island (13 from 46-0 to 92-0 mm.), 
and Sherm Sheikh (9 from 20-0 to 74-0 mm.). 

This species is apparently confined to the Red Sea. 



Sub-order SCLEROPAREI 

SCORPAENIDAE 

*Scorpaenopsis albobrunneus (Gunther) 

Twenty-two specimens, all obtained from pieces of coral brought up by a diver 
at the following localities: Dahab (8 from 19-0 to 44-0 mm.), Tiran Island (2 of 35-0 
and 40-0 mm.), Sanafir Island (7 from 21 -o to 48-0 mm.), Sherm Sheikh (5 from 19-0 
to 35-0 mm.). 

Pterois volitans Linnaeus 

Three specimens taken at Dahab (1 of 155 mm.), Hobeik (1 of 190-0 mm.), and Abu 
Zabad (i of 51-0 mm.). 

Order DISCOCEPHALI 

ECHENEIDIDAE 

Echeneis neucrates Linnaeus 
One specimen of 617 mm. caught by hand-line at Sherm Sheikh. 

Order PLECTOGNATHI 

Balistidae 

Odonus niger (Ruppell) 

Six specimens caught by hand-line at Sherm Sheikh (4 from 143-0 to 186-0 mm.) 
and Hobeik (2 of 240-0 and 285-0 mm.). 

Balistapus undulatus (Mungo Park) 

Two specimens. One of 184 mm. caught by hand-line at Hobeik and one of 35 mm. 
obtained from a piece of coral. 

Rhinecanthus assasi (Forskal) 

Four specimens taken at Abu Zabad (2 of 205 and 210 mm.) and Sanafir Island 
(2 of 170 and 190 mm.). 

This species seems to be restricted to the Red Sea, the Gulf of Aden, and the 
Indian Ocean coast of Arabia. 



244 THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 

Aluteridae 

Oxymonacanthus halli sp. nov. 
(Fig. 3) 

Two specimens, the holotype of 38-0 mm. and one paratype of 39-5 mm. taken 
from a piece of coral at Sanafir Island in the northern Red Sea. 

(In the description which follows, measurements and counts of the holotype 
precede those of the paratype which are placed in brackets.) 

Body proportions (relative to a standard length of 100) : Greatest depth 35-5 (36-1) ; 
length of head 34-8 (35*4) ; predorsal length (from tip of snout to origin of dorsal fin) 
55*5 (55*7) i preanal length 6i-8 (62-0) ; depth of caudal peduncle 14-4 (13*3) ; length 
of pectoral fin 9-9 (9-5) ; height of first dorsal spine 23-0 (24-0). 




Fig. 3. Oxymonacanthus halli sp. nov. 

Head proportions (relative to a head length of 100) : Length of snout 64-0 (64*3) ; 
horizontal diameter of eye 26-4 (25-0). 

Fin rays: Dorsal II, 28 (II, 27) ; anal 27 (25) ; left pectoral 10 (10) ; caudal 12. 

Body covered with very numerous spinules, which become larger (about twice the 
length of those immediately behind the eye) and fewer on the caudal peduncle, 
particularly over the lateral, median regions. These spinules extend out to about 
three-quarters the length of each caudal ray and are not found on the tip of the snout 
in front of the brown pigment band which encircles it. First dorsal spine studded 
anteriorly and laterally with blunt spines, while posteriorly there are two rows of 
9 or 10 rather larger blunt spines, the lower of which, at least, project backwards and 
downwards. Second dorsal spine very small. Immediately posterior to the first 
dorsal spine there is a fairly deep, wide groove in the back, of much the same length 
as this spine. 1 Pelvic spine supporting a small ventral flap. Dorsal, anal, and 
pectoral rays unbranched. Caudal rays branched except for the upper and lower 
outermost rays which are stouter at the base than the inner rays. Jaws meeting 
dorsally at the tip of the snout. 

General colour blue with longitudinal rows of roughly circular deep yellow spots. 

1 Presumably the long dorsal spine folds into this groove, but I have been unable to unlock the trigger 
mechanism by pressure on the small second spine. 



THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 245 

Between the origin of the dorsal and anal fins nine rows of these spots can be counted. 
Tip of snout yellow in front of the brown pigment ring which encircles it. Membrane 
of dorsal fin yellow, pelvic flap orange with a black edging. Caudal with a black 
vertical bar of pigment. Iris golden with six symmetrically arranged slate blue 
sectors. 

This species differs from Oxymonacanthus longirostris (Bloch & Schneider), the only 
other species of this genus, in the following : 





0. halli 


0. longirostris 1 


Dorsal rays . 


27 and 28 


31-32 


Anal rays . 


25 and 27 


29-30 


Pectoral rays 


10 


11-12 



In addition there are certain differences in the colour pattern. 

1. In Oxymonacanthus halli there are no longitudinal yellow stripes in front of the 
eye as are usually found in 0. longirostris. 

2. There are 9 longitudinal rows of yellow spots (counting across the body between 
the origins of the dorsal and anal fins) in 0. halli, whereas in longirostris there 
are usually 7 (occasionally 6 or 8). The number of spots in each row is also 
greater in the new species. There are 18 or 19 (counting along the row behind 
the eye), whereas in longirostris there are 12-16. 

3. In 0. longirostris there is usually a small area of the abdomen just above the 
pelvic flap which is differentiated from the rest of the body by being brown 
in colour and dotted with small white spots. This is absent in the two specimens 
of 0. halli. 

I have much pleasure in naming this species after Major H. W. Hall, M.C., the 
owner of M.Y. Manihine. 

OSTRACIONTIDAE 

Ostracion tuberculatus Linnaeus 
Two specimens of 270 and 300 mm. taken by cast-net at Sanafir Island. 

Lagocephalidae 

Lagocephalus sceleratus (Forster) 

Four specimens from 260 to 280 mm. taken by hand-line at a depth of 10 fathoms 
at Sanafir Island. 

Tetraodontidae 

Amblyrhynchotes diadematus (Ruppell) 

One specimen of 146 mm. taken by cast-net at Mualla. This species is confined 
to the Red Sea. 

1 Counts and measurements based on specimens from Samoa (1) (standard length 79-0 mm.), Am- 
boyna (1 of 70-0 mm.), Ponape, Caroline Islands (2 of 43 and 65 mm.), New Britain (1 of 57-0 mm.), and 
one (no locality given) from Bleeker's collection (67 mm.). 



246 THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 

Arothron hispidus (Linnaeus) 

Two specimens of 285 and 340 mm. taken by cast-net at Sanafir Island. 

These two specimens and another from the Gulf of Suez all have much more 
numerous and smaller white spots on the body than in examples taken outside the 
Red Sea. 

Canthigasteridae 

*Canthigaster cinctus (Solander) 
One specimen of 65 mm. taken by a fish-trap at Aqaba from a depth of 10 fathoms. 

DISCUSSION 

Among the 113 species considered in this report are a number which appear to be 
confined to the Red Sea. They may be subdivided as follows : 

A. Almost certainly endemic 

Pseudochromis olivaceus Ruppell 

Crenidens crenidens crenidens (Forskal) 

Diplodus noct ((Ehrenberg) Cuvier & Valenciennes) 

Chaetodon fasciatus Forskal 

Dascyllus marginatus marginatus (Ruppell) 

Haliophis guttatus (Forskal) 

Hypoatherina gobio (Klunzinger) 

Amblyrhynchotes diadematus (Ruppell) 

B. Possibly endemic 

Hetereleotris vulgare Klunzinger 
Enchelyurus kraussii Klunzinger 
Oxymonacanthus halli Marshall 

(The first two species are small and inconspicuous) 

C. Species with Red Sea forms distinguishable from those of the Indian Ocean 

Spratelloides gracilis (Schlegel) 
Therapon jarbua (Forskal) 
Platax orbicularis (Forskal) 
Arothron hispidus (Linnaeus) 

The number of species collected by this expedition probably represents about 
one-fifth of the total fish fauna of the Red Sea (Klunzinger, 1870 and 1871, lists 
about 490 species). If it is a representative sample, then about 10 per cent, of the 
species (and sub-species) known from this area are endemic. Moreover, to judge 
from the work on this collection, this percentage may well prove to be considerably 
higher, when more material becomes available for study. 

Before discussing how these endemic elements may have originated it will be 
necessary to outline briefly the geological history of the Red Sea area. Although 



THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 247 

the evidence is rather incomplete it seems that the formation of the main physical 
features were completed during the Pliocene and that during this time the Red Sea 
became connected with the Gulf of Aden and the Indian Ocean. Fox (1926) suggests 
that the invasion of Indian Ocean species into the Red Sea occurred some time after 
the Middle Pliocene. (Earlier an ancestral Red Sea appears to have come into being 
as the result of the faulting of Eocene strata followed by the filling of the resulting 
depression with water from the north. Later on the Red Sea appears to have lost 
its connexion with the northern Tethys Sea, for during Miocene times it shrank in 
area giving rise to great deposits of rock salt.) Continuing from middle Pliocene 
times there seems no doubt that there was again a connexion between the Medi- 
terranean (Tethys Sea) and the Red Sea (the latter now containing a mixed 
Mediterranean and Indian Ocean fauna), but when the Gulf of Suez became cut off 
from the Mediterranean is not very certain. 

This would seem to be the generally accepted geological history of the Red Sea, 
but Sewell (1948) has considered the implications of Zeuner's (1945) work on the 
Pleistocene period. Zeuner suggests that during the last Glacial period the sea-level 
fell as the result of ice formation, his figure for the Mediterranean being —100 
metres, while a low level of —200 metres has been suggested for the penultimate 
glaciation. 

Sewell (1948) concludes that this lowering of sea-level might well have left the 
shallow sill at the southern end of the Red Sea uncovered, which ' . . . must have 
resulted in the almost complete disappearance of the Red Sea as it exists today and 
its reduction to two small inland lakes which were in all probability hypersaline. 
Under such changes as these it is difficult to suppose that anything of the marine 
fauna can possibly have survived, and the original fauna of the Tethys Sea that was 
derived from the Indo-Pacific region must have disappeared.' Following from this 
the sea-level once again rose at the end of the Glacial period, resulting in a second 
and final influx of species from the Indian Ocean into the Red Sea. 

Concerning the origin of forms peculiar to the Red Sea there are certainly two 
possibilities : 

1. That they have evolved from species entering the Red Sea. 

2. That they may be the only survivors of species which originally lived in the 
Indian Ocean. It might be suggested, for example, that the Indian Ocean 
representatives of these species have been eliminated during geological history 
whereas conditions in the Red Sea favoured their survival. 

A third possibility of whether the endemic forms are relics from the ancestral Red 
Sea seems so unlikely that it will not be considered beyond pointing out that the 
presence of great rock-salt deposits, probably of Miocene age, implies that this early 
sea must have been subject to extensive evaporation. As already mentioned, Sewell 
(1948) concluded that this would be likely to happen and that it most probably 
resulted in a mass extinction of the marine fauna. 

Beginning with the second suggestion, it seems somewhat improbable that this 
fairly high number of endemic forms should have all possessed the potentialities of 
surviving in the Red Sea while the Indian Ocean ancestral stock perished. Following 

zoo. 1.8. Kk 



248 THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 

the formation of the 'modern' Red Sea the main hydrological features would 
gradually have evolved, that is, higher summer temperatures and greater salinities, 
which now distinguish it from the Indian Ocean. Such changes would have tended 
to bring about correlated changes in the fish fauna (among them extinction) rather 
than the preservation of species. To put it another way, it seems unlikely that these 
forms should have all been pre-adapted to conditions in the Red Sea. While it is not 
possible to state the latter with certainty, it is of interest that the Red Sea supports 
fewer species of fishes than the Indian Ocean. Sewell (1948) has similar findings for 
the free-swimming planktonic Copepoda and suggests that many species which are 
widely distributed in the Indo-Pacific are unable to survive the changes in salinity 
and temperature on being carried into the Red Sea. 

Turning to the first suggestion, if a number of the ancestral Indian Ocean immi- 
grants have evolved into species and sub-species peculiar to the Red Sea, there should 
be some evidence for this today. It would be reasonable to expect to find at least 
some of these endemic Red Sea forms pairing off with the present-day Indian Ocean 
representatives. Regan (1906-1908) and Meek & Hildebrand (1923), when consider- 
ing the marine fishes of Panama, have remarked on the many close parallels between 
the faunas of the opposite sides of the isthmus. It is generally considered that the 
formation of the Isthmus of Panama during late Pliocene times separated many 
species into Atlantic and Pacific populations which have diverged in isolation. 

In the Red Sea there are certain endemic species which are paired with others from 
the Indian Ocean. From this collection there are the following pairs: 

Red Sea Indian Ocean 

Diplodus nod (C.V.) Diplodus sargus (L.) (also occurs in the Med.) 

Chaetodon fasciatus Forskal Chaetodon lunula Lacepede 

Haliophis guttatus Forskal Haliophis malayanus M. Weber 

Oxymonacanthus halli Marshall Oxymonacanthus longirostris (Bloch & Schneider) 

While the members of these pairs may well have arisen by the separation of an 
original species into Red Sea and Indian Ocean populations, they are not sufficiently 
closely related to draw any certain conclusions as to their past history. Instead, it 
will be better to concentrate on infra-specific categories. Here there are the proposed 
sub-species of Dascyllus marginatus and the examples listed earlier of differences 
between Red Sea and Indian Ocean populations of certain species. Judging from the 
impressions gained in working out this collection and from numerous instances in 
the literature 1 where Red Sea examples of a species can be distinguished from others 
from the Indian Ocean, there can be little doubt that when good series of specimens 
from both areas are available, more species will be found to have Red Sea 'forms'. 

The evolution of these endemic elements implies that after entering the Red 
Sea they became isolated to some degree. Leaving aside problems concerning the 
Suez Canal, entry via the Gulf of Aden is through the narrow Strait of Bab-el Mandeb, 
inside which is a shallow sill, where the greatest depth is only about 100 metres. 
Climatic conditions and the basin-like character of the Red Sea are the predominating 

1 Particular reference may be made to Fraser-Brunner (1950), who remarks that ' . . . among the Chaeto- 
donts at least I find that few or none of the known Red Sea forms are identical with those outside'. 



THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 249 

factors controlling the temperature and salinity of the waters, and as already men- 
tioned, the latter features were evolved after the formation of the ' modern ' Red Sea. 
Today very soon after entering the Red Sea the salinity rises by about 2% and in 
summer the surface temperature increases by about 3-5 ° C. In winter there appears 
to be little difference between the surface temperatures of the Red Sea and the Gulf 
of Aden (data from Sverdrup, Johnson, & Fleming, 1942). 

Perhaps this quite abrupt change in one or both physical factors may be a barrier 
to the exchange of Red Sea and Gulf of Aden fishes and has been so long enough for 
new forms to have arisen. Perhaps the habits of the fishes themselves may be another 
factor, species which are closely dependent on coral life and less migratory being more 
prone to differentiation than the more active pelagic species. (While there is the 
possibility of the larvae of the former types being carried out of the Red Sea (or 
into it), younger stages are usually more 'exacting' than adults in the physical con- 
ditions necessary for their existence; thus such an event may prove disastrous.) 
Again owing to the changes in temperature and salinity which have occurred since 
the formation of the Red Sea, certain species may have become reproductively iso- 
lated from their Indian Ocean ancestors, through the evolution of differing breeding 
seasons. In conclusion, however, it should be added that these are no more than 
suggestions to be tested in the light of further knowledge. 

If more data were available on the fish fauna it would be interesting to compare 
and contrast the Red Sea-Indian Ocean relationships with those found across the 
Straits of Panama. Concerning the latter area, Gilbert & Starks (1904) in discussing 
the parallels between the two faunas concluded that : 

' The ichthyological evidence is overwhelmingly in favour of the existence of a former open 
communication between the two oceans, which must have closed at a period sufficiently remote 
from the present to have permitted the specific differentiation of a very large majority of the 
forms involved. That this differentiation progressed at widely varying rates in different in- 
stances becomes at once apparent. A small minority of the species remain wholly unchanged, so 
far as we have been able to determine that point. A large number have become distinguished 
from their representatives of the opposite coast by minute (but not "trivial") differences which 
are wholly constant. From such "representative forms" we pass by imperceptible gradation to 
species much more widely separated whose immediate relation in the past we cannot confidently 
affirm.' 

Later work by Meek & Hildebrand (1923) did not change these conceptions, except 
that it was found that fewer species could properly be regarded as common to both 
coasts and more species were discovered with representative Atlantic and Pacific 
forms. 

It is not proposed on the present limited data to draw conclusions regarding rates 
of evolution in the Red Sea fauna. Direct comparison with the Panama findings is 
not, of course, possible for two main reasons: firstly that there is a connexion between 
the Red Sea and the Indian Ocean (which may make for genetic interchange between 
the two faunas), and secondly, that there are often greater differences in temperature 
(but not in salinity) between Red Sea and Indian Ocean waters than exist across the 
Straits of Panama (this aspect will be discussed later). Whether the degree of 
endemism of the Red Sea fauna could have been attained since the last Glacial 



250 THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 

period (if Zeuner's (1945) figures of drop in sea-levels and Sewell's (1948) conclusions 
from these are considered), is a question which will best be considered when the 
large collection of fishes recently obtained from Sudanese waters has been studied. 

Finally the fact that the Red Sea is for part of the year warmer and always more 
saline than Indian Ocean waters must be considered as a ' conditioning factor ' in the 
evolution of Red Sea forms. Before this can be done a list of the differences between 
closely related forms will be given. 

Spratelloides gracilis. The Red Sea populations tend to have fewer pectoral and 
anal rays than those from the Japanese area. 

Therapon jarbua. The Red Sea form has fewer dorsal spines and a slimmer body 
form. 

Diplodus nod. This differs from D. sargus from the Indian Ocean in the slimmer 
body form, the tendency for the dorsal and anal fins to have fewer rays, the smaller 
number of scale rows above and below the lateral line, and the fewer gill-rakers on 
the first arch. 

Crenidens crenidens. The Red Sea sub-species differs in the slimmer body form, the 
fewer scale rows above and below the lateral line, and the lesser relative height 
and length of the soft dorsal and pelvic fins respectively. 

Platax orbicularis. Red Sea examples tend to have fewer pectoral rays than those 
from the Indo-Pacific. 

Dascyllus marginatus. The Red Sea sub-species differs from that of the Indian Ocean 
in the tendency to have fewer pectoral rays, relatively longer soft dorsal and anal 
fins, and generally lesser developed pigmentation. 

Oxymonacanthus halli. Differs from 0. longirostris from the Indo-Pacific in having 
fewer dorsal, anal, and pectoral rays. There are also differences in the colour pattern. 

It is interesting to consider these differences in relation to present data concerning 
the correlations of character with environment in fishes. It is well known that the 
number of fin rays and scales often appears to be inversely related to the temperature 
with which the above data appear to be in agreement. But a study of the charts of 
surface temperatures contained in the Monthly Meteorological Charts of the Indian 
Ocean (M.O. 519. H.M. Stationery Office) shows that from January until May 
northern Red Sea waters are consistently cooler than those of the Indian Ocean, 
while evidence is accumulating that many Red Sea fishes spawn during January and 
February — a problem to be discussed more fully in a later paper. On the other hand, 
the number of parts of a fish may have a direct relationship with salinity. Precisely 
what would be the apparent effect of high temperatures and increased salinities on 
numbers of fin rays or scales in Red Sea fishes (compared to their nearest relatives 
from the Indian Ocean) is impossible to predict. However, recent work by Heuts 
(1949) showing the combined effect of temperature and salinity on the number of fin 
rays in Gasterosteus aculeatus may be of particular significance here. Considering only 
the marine B population, increase in salinity at io° C. led to an increase in the mean 
number of dorsal and anal rays, whereas at 23 C. the effect of this was to produce a 
decrease. 

Concerning body form, Hubbs (1940) states that : ' Forms of warmer water, and in 
the sea those of brackish water, typically have deeper bodies and larger heads than 



THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 251 

those of colder or more saline waters. ' In the Red Sea there may be a correlation with 
the increased salinity for in three of the examples listed above, the Red Sea form had 
a slimmer body shape. More data are desirable before arriving at any conclusions, 
but these are interesting problems and further comparative studies of Red Sea and 
Indian Ocean fishes may well contribute to a closer understanding of them. At the 
same time experimental studies would be desirable. It need only be added that close 
correlation between environment and structure need not mean that the changes are 
entirely dependent on the environment. There is evidence from other work that the 
genotype is also involved. It will be apparent from the recognition of certain sub- 
species and the trend of this discussion that the view is held that it is unlikely that 
these correlations solely arise from the action of the environment on the phenotype. 

To conclude, it looks as though partially enclosed seas, such as the Red Sea, may 
be centres for the evolution of new forms. I am much indebted to Dr. A. E. Parr for 
drawing my attention to the Gulf of California, which also harbours certain endemic 
species and sub-species. Setchell (1937), referring to earlier work in the gulf, points 
out that fifteen species or varieties of Sargassum are endemic, '. . . thus indicating 
what is borne out by the remaining marine flora of this body of water, that it forms a 
"pocket" of more than ordinary distributional interest'. Burkenroad (1938) notes 
that certain penaeid prawns are confined to the Gulf of California, while Parr (193 1) 
took certain species of deep-sea fishes in the gulf that had originally been described 
from there by Garman (1899) and have not so far been taken outside this area 
(although neighbouring areas have been well worked). 

More hydrological and biological data will be necessary to discover the degree 
of isolation of the fauna of these marine pockets. Mayr (1942) has remarked that: 
'In the sea isolation is rarely complete and the partially isolated populations are 
normally very large. It is mainly for this reason that marine species have fewer sub- 
species than terrestrial species and that the entire evolution in the sea is slower and 
more conservative.' Further work on partially enclosed areas should help towards 
an understanding of the evolution of new forms in the seas. 



REFERENCES 

Breder, C. M. 1938. A contribution to the life histories of Atlantic Ocean Flying fishes. Bull. 

Bingham oceanogr. Coll. 6 (5) : 1-48. 
Brunner, A. F. 1950. Holacanthus xanthotis, sp.n., and other chaetodont fishes from the Gulf 

of Aden. Proc. zool. Soc. Lond. 120: 43-48. 
Bruun, A. F. 1935. Flying fishes (Exococtidae) of the Atlantic. Systematic and biological 

studies. Dana Rep., Copenhagen, 6: 1-108. 
Burkenroad, M. D. 1938. The Templeton Crocker Expedition XIII. Penaeidae from the 

region of Lower California and Clarion Island, with descriptions of four new species. Zoo- 

logica, N.Y. 23: 55-91. 
Duncker, G., & Mohr, E. 1925. Die Fische der Siidsee-Expedition der Hamburgischen Wissen- 

schaftlichen Stiftung 1908-1909. 1 Teil (Fistulariidae, Centriscidae, Syngnathidae) . Mitt. 

zool. Stlnst. Hamburg, 41: 93-112. 
Fowler, H. W. 1933. Contributions to the biology of the Philippine Archipelago and adjacent 

regions. The fishes of the families Banjosidae, Lethrinidae, Sparidae, Girellidae, Kyphosidae, 

Oplegnathidae, Gerridae, Mullidae, Emmelichthyidae, Sciaenidae, Sillaginidae, Arripidae, 



252 



THE 'MANIHINE' EXPEDITION TO THE GULF OF AQABA 



and Enoplosidae collected by the United States Bureau of Fisheries Steamer Albatross 

chiefly in Philippine Seas and adjacent waters. Bull. U.S. nat. Mus. No. ioo (12) : 1-465. 
Fox, H. M. 1926. Zoological results of the Cambridge Expedition to the Suez Canal, 1924. 

Trans, zool. Soc. Lond. 22: 1-64. 
Garman, S. 1899. Reports on an exploration off the west coasts of Mexico, Central and South 

America, and off the Galapagos Islands in charge of Alexander Agassiz, by the U.S. Fisheries 

Commission Steamer Albatross during 1891, Lieut. Commander Z. L. Tanner, U.S.N. 

Commanding. The Fishes. Mem. Harv. Mus. comp. Zool. 24: 1-431. 
Gilbert, C. H., & Starks, E. C. 1904. The fishes of Panama Bay. Mem. Calif. Acad. Sci. 

4: 1-304- 
Heuts, M. J. 1949. Racial divergence in fin ray variation patterns in Gasterosteus aculeatus. 

J. Genet. 49: 1 83-191. 
Hubbs, C. L. 1940. Speciation of fishes. Amer. Nat. 74: 198-21 1. 
Jordan, D. S., & Thompson, W. F. 191 2. A review of the Sparidae and related families found 

in the waters of Japan. Proc. U.S. nat. Mus. 41: 521-601. 
Klunzinger, C. B. 1870. Synopsis der Fische des Rothen Meeres, Part I. Verh. zool. bot. Ges. 

Wien, 20: 669-834. 

1871. Part II. Verh. zool. bot. Ges. Wien, 21: 441-688. 

1884. Die Fische des Rothen Meeres. 1. Theil. Acanthopteri veri, Owen. 1-133. Stuttgart. 

Mayr, E. 1942. Systematics and the origin of species: xiv + 334 pp. New York. 

Meek, S. E., & Hildebrand, S. F. 1923. The marine fishes of Panama, Part I. Field Mus. 

Publ. Zool. 15: 1-330. 
Parr, A. E. 193 1. Deep sea fishes from off the western coast of North and Central America. 

Scientific Results of the Second Oceanographic Expedition of the Pawnee 1926. Bull. 

Bingham, oceanogr. Coll. 2 (4) : 1-53. 
Regan, C. T. 1906-1908. Biol. Cent.-Amer. Pisces: 1-192. 

Ruppell, E. 1828. Fische des Rothen Meeres. Atlas Reise nordl. Afrika, Part IV: 1-144. 
Schultz, L. P. 1943. Fishes of the Phoenix and Samoan Islands collected in 1939 during the 

expedition of the U.S.S. Bushnell. Bull. U.S. Nat. Mus. 180: x + 316. 
Setchell, W. A. 1937. The Templeton Crocker Expedition of the California Academy of 

Sciences 1932, No. 34. Report on the Sargassums. Proc. Calif. Acad. Sci. 22: 127-158. 
Sewell, R, B. S. 1948. The Free-swimming planktonic Copepoda: Geographical Distribution. 

/. Murray Exped. 1933-34. Sci. Rep. 8: 317-592. 
Smith, J. L. B. 1938. The South African fishes of the families Sparidae and Denticidae. Trans. 

roy. Soc. S. Afr. 26: 225-305. 

1949. The Sea Fishes of Southern Africa: xvi + 550 pp. Cape Town. 

Sverdrup, H. U., Johnson, M. W., & Fleming, R. H. 1942. The Oceans: x + 1087 pp. New 

York. 
Weber, M., & de Beaufort, L. F. 1929. The Fishes of the Indo- Australian Archipelago, 5: 

xiv + 458pp. Leiden. 

193 1. Ibid. 6: xii + 448 pp. Leiden. 

1936. Ibid. 7: xvi + 607 pp. Leiden. 

Zeuner, F. E. 1945. The Pleistocene Period: its climate, chronology and faunal successions: 

xii-l-322 pp. Ray Soc. London. 






1352 




PRINTED IN 

GREAT BRITAIN 

AT THE 

UNIVERSITY PRESS 

OXFORD 

BY 

CHARLES BATEY 

PRINTER 

TO THE 

UNIVERSITY 



ON THE SPECIES AND 

RACES OF 

THE YELLOW WAGTAILS 

FROM WESTERN EUROPE 

TO WESTERN NORTH 

AMERICA 

C. H. B. GRANT 

and 

C. W. MACKWORTH-PRAED 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ZOOLOGY Vol. i No. 9 

LONDON : 1952 



ON THE SPECIES AND RACES OF 

THE YELLOW WAGTAILS 

FROM WESTERN EUROPE TO 

WESTERN NORTH AMERICA 



BY 



C. H. B. GRANT 



and 






C. W. MACKWORTH-PRAED 







Pp. 253-268; Pis. 33-35 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ZOOLOGY Vol. 1 No. 9 

LONDON : 1952 



THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted in 1949, is issued 
in five series, corresponding to the Departments of the 
Museum. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four hundred 
pages, and will not necessarily be completed within one 
calendar year. 

This paper is Vol. 1, No. 9 of the Zoology series. 



PRINTED BY ORDER OF THE TRUSTEES OF 
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Issued July 1952 Price Five shillings 



Bull. 8. M. (N. H.) Zoology, I, 9 



PLATE 33 














Drawn by H. Gronvold and C. E. Talbot Kelly. 



Blue-headeci Yellow-Wagtail, male 
Budytes flavus flavus 

Budytes flavus dombrowskii, male 

Yellow Wagtail, male 
Budytes luteus luteus 

Budytes luteus flavissima, male 

Dark-headed Yellow-Wagtail, young 
Budytes thunbergi thunbergi 

Black-headed Yellow-Wagtail, young 
Budytes feldegg 



Blue-headed Yellow-Wagtail, female 
Budytes flavus flavus 

Budytes flavus dombrowskii, female 

Yellow Wagtail, female 
Budytes luteus luteus 

Dark-headed Yellow-Wagtail, male 
Budytes thunbergi thunbergi 

Black-headed Yellow-Wagtail, male 
Budytes feldegg 

White-browed Yellow-Wagtail 
Budytes superciliaris 



Blue-headed Yellow-Wagtail, young 
Budytes flavus flavus 

Yellow-browed Yellow-Wagtail, male 
Budytes perconfusus 

Yellow Wagtail, young 
Budytes luteus luteus 

Dark-headed Yellow-Wagtail, female 
Budytes thunbergi thunbergi 

Black-headed Yellow-Wagtail, female 
Budytes feldegg 

White-headed Yellow-Wagtail, male 
Budytes leucocephalus 



ON THE SPECIES AND RACES OF THE 

YELLOW WAGTAILS FROM WESTERN EUROPE 

TO WESTERN NORTH AMERICA 

By c. h. b. grant and C. W. MACKWORTH-PRAED 

{Received 15.V.51) 
SYNOPSIS 

An endeavour has been made to collect all the known relevant facts on this group and to show that it 
is not correct to place all the Yellow Wagtails in one species. The authors have based their main con- 
clusions on adult males, and the measurements given are only of adult males from the breeding area, so 
that a true comparison can be given, and there can be no confusion with measurements of specimens from 
the non-breeding areas which may have been misidentified by us. It is perhaps of interest to note that the 
southern species have usually a white chin and throat and the Far Eastern tend to have a longer hind claw. 

The normal migration route appears to be mainly north and south, although there is largely a tendency 
to a north-eastern to south-western movement. 

Measurements appear to be of little value in determining the species and races except perhaps in the 
case of the hind claw of B. thunbergi macronyx. 

There is still much to learn about this group, especially the exact breeding- ranges of the species and 
races. 

It should be remarked that adult males of all species and races are inclined to have some olivaceous 
green on the crown, or yellow in B. leucocephalus, and a broken spotted collar in both sexes. These are 
not specific or racial characters and may be individual retention of juvenile plumages. 

A total of 2,594 specimens have been examined. 

Many authors 1 have written on this group, adding considerably to our general 
knowledge, and several have described new races. We have lately had occasion to 
study these Wagtails critically, with especial regard to those species and races which 
occur in Africa during the non-breeding season. We found it necessary, however, to 
survey the whole group. 

The usual English practice has been to place all as races of one species, Budytes 
flavus (Linnaeus), but several authors have divided them into a number of species. 
We have examined all the literature we can find, the large series of specimens in the 
British Museum collection, and had the kind loan of specimens from Colonel Meinertz- 
hagen, Colonel Payn, Major Payn, M. Mayaud, Dr. K. H. Voous, the Copenhagen 
Museum, the South African Museum, the Royal Natural History Museum, Stock- 
holm, and the Coryndon Museum, Nairobi. 

We are of the opinion that the genus Budytes should be retained as these Yellow 
Wagtails have somewhat different habits to the Black and White Wagtails, and 
behave more like Pipits in many respects. It is very unfortunate that we are so out 
of touch with the Russian museums, as no doubt they have series of birds from the 
breeding areas which would have been most valuable to examine, but no doubt a 
number of them have been recorded in the Russian journals we have consulted. 

The maps we give not only show the known breeding-areas and limits of movements 
in the non-breeding season, but also the comparatively vast areas in Europe and 

1 The bibliography at the end of this article covers the principal references on this group. 



256 ON THE SPECIES AND RACES OF THE YELLOW WAGTAILS 

Asia from which no breeding birds have been recorded, and surely some of these — 
especially along the rivers — must hold ground suitable to the Yellow Wagtail, 
although Sushkin (1925) states that apparently none are found breeding in the lower 
part of the Kobdo basin. 

Our long and very careful examination convinces us that those authors who 
recognize several species are correct, and the maps we give show that there is in 
several cases an overlap in breeding distribution, a fact supporting the recognition of 
species. We have divided this group into seven species and would remark that 
Budytes flavus becomes paler on the head as it goes eastward, the palest being a 
specimen from Lake Aral, and where the breeding area of this race and B. f. beema 
meet specimens show characters of both. We feel sure that Budytes thunbergi, B. 
luteus, and B. feldegg should be treated as species and that B. superciliaris and B. 
leucocephalus are also recognizable as distinct species. In the course of this examina- 
tion we have decided to name a new race and a new species, one from Lake Alakul 
on the Mongolian border, west of Dzungaria, on a single male that does not fit in 
with any other Yellow Wagtail without an eyestripe, and the other from five adult 
male specimens that are all so exactly alike and with such distinct characteristics 
that it would be unbelievable they do not represent an undescribed species. On the 
original labels of two from Khartoum A. L. Butler recorded his opinion that they 
are 'possibly hybrids M.f. rayi and M.flava', and on the male from the Copenhagen 
Museum ' rayixflaval ', but we do not think that this is so, as their characteristics 
do not fit in with what would be expected of such an intermediate and they are all 
exactly alike. 

The Yellow Wagtails have been credited with being a very variable group and 
any specimen not fitting into the general rule was merely passed over as an aberrant. 
This we consider a mistaken and dangerous point of view liable to obscure completely 
the true picture. Individual variation there is, but within the species. We do not 
find this group particularly difficult to disentangle and we advance no theories about 
it (Johansen, 1946), having based our conclusions on the facts as shown by specimens 
and the recorded known breeding distribution. Young birds in their first plumage in 
all the species are more or less ashy above with dusky centres to the feathers; a 
blackish streak between the crown and light eyestripe ; below more or less buffish 
white with a black moustachial stripe joining up to an almost perfect collar on lower 
neck. In B. feldegg the dusky centres to the feathers of the upper parts are broader 
and darker, and in the B. luteus group these markings are almost absent and there 
is a tendency to a yellow wash on the lower belly. In this dress they can be named 
from the local population in which they occur, as they do not apparently leave their 
breeding grounds until they have moulted into an immature (intermediate) dress. 
In this immature dress they are found in their non-breeding quarters, and as the 
species and races may occur in mixed flocks it is not easy to name them correctly. 

Comparison with correctly identified adult females does reveal certain similarities 
by which the majority can be named, but no written description can give those small 
differences which the eye can spot when the group as a whole is closely and meticu- 
lously studied. Even so, every immature specimen can by no means be named with 
absolute certainty. 



FROM WESTERN EUROPE TO WESTERN NORTH AMERICA 257 

It has been said that there is a difference in length of tail between B. flavus flavus 
and B. flavus beema and other named races, but we have measured birds from the 
breeding areas and cannot agree that this is so. 

The seven species which we recognize can be distinguished as follows : 

A . A streak from base of bill to over and behind eye in male : 

Budytes flavus (Linnaeus). Blue-headed Yellow Wagtail 
Budytes flavus flavus (Linnaeus) 

Motacilla flava Linnaeus, 1758, Syst. Nat. 10th ed. : 185, South Sweden. (For synonyms see 
Hartert, 1905, Vog. Pal. Fauna: 287.) 

Adult male. A distinct white streak from base of bill to over and behind eye, very 
rarely indeed broken over the eye ; head and neck and sides of face grey (variable 
individually) ; usually flecked with white on ear-coverts ; usually some white on chin ; 
rest of underparts bright yellow; sometimes some spots on lower neck. Wing 77-85, 
hind claw 6-11, tail 67-74 mm - Twenty-one males from breeding area measured, 
a total of 474 specimens examined. 

The female has the head and neck more olivaceous ; below, chin and neck bufnsh 
white ; rest of underparts pale yellow ; often with spots on lower neck. 

Distribution: Breeding southern Norway, southern Sweden, southern Finland, 
eastern England (rarely) to northern, western, and central France, middle Europe 
and the Caspian Sea ; in non-breeding season to Africa throughout and Arabia. 

Mayaud, 1949, states that at Oleron, western France, intermediates occur between 
B. flavus and B. fasciatus. Through the kindness of Dr. Mayaud we have seen three 
breeding males, Mayaud Nos. 2333, 2334, and 2335, all taken in May, and consider 
them to be B. f. flavus. 

In British Birds: 86, 1949, Stuart Smith and Ramsden record a variant yellow 
Wagtail breeding in the hills near Higher Disley in Cheshire in June. They do not 
quite agree as to the exact markings of the head of the male and, anyway, such sight 
records are most difhcuTt to fix and are often not worthy of being recorded. It would 
appear that this is possibly another record of Budytes f. flavus breeding in England. 

Budytes flavus beema Sykes 

Budytes beema Sykes, 1832, Proc. zool. Soc. Lond.: 90, Deccan, India, of which Budytes dubius 
vel anthoides Hodgson, 1844, in Gray's Zool. Misc. : 83 (nom. nuda) and Budytes brevicaudatus 
Homeyer, 1878, /. Om. Lpz., 131, Etawah, north-western India, are synonyms. 

Adult male : Head and neck pale french grey, variably darker or paler. In freshly 
moulted dress the mantle is lighter and yellower than in the breeding season. Wing 
76-81, hind claw 9-10, tail 67-72 mm. Nine males from breeding area measured; a 
total of 320 specimens examined. Where this race meets the nominate race specimens 
may be placed in either. The female is not distinguishable from the nominate race. 

Distribution : Breeding from the Ural Mts. and Caspian Sea to Tomsk and Turkes- 
tan ; in non-breeding season to the Sudan, Kenya Colony, Nyasaland, Arabia, and 



258 ON THE SPECIES AND RACES OF THE YELLOW WAGTAILS 

India. Main non-breeding quarters appears to be India. One specimen from 
Valencia, Spain in April. 

Budytes flavus fasciatus Zander 

Budytes fasciatus Zander, 1851, Naumannia 1 (4): 19, southern France, of which Motacilla 
flava iberiae Hartert, 1921, Vog. Pal. Fauna 3: 2097, southern France, is a synonym, but if 
these Yellow Wagtails are placed in the genus Motacilla the latter name must be used as 
Motacilla fasciata (Zander) is preoccupied by Motacilla fasciata Bechstein. 

Adult male : White streak from base of bill to over and behind eye ; chin to neck in 
front white. Wing 75-82, hind claw 8-11, tail 67-72 mm. Fifty-seven males from 
breeding area measured, a total of ninety-six specimens examined. The female also 
has the chin to neck in front white ; the head is duller grey and below, pale or buffish 
yellow, often with spots forming a sort of collar on the lower neck. In fresh dress the 
head is more olivaceous. 

Distribution: Breeding Spain, Portugal, eastern Pyrenees to western areas of 
southern France as far east as the Camargue, the Balearic Islands, and Morocco ; in 
non-breeding season to Italy, Morocco, Algeria, Tunisia, and French Sudan. 

Wardlaw Ramsay, 1923 (Birds of Europe and North-west Africa: 61) states that this 
race breeds in Algeria, and this has been quoted by other authors. We cannot find 
any evidence in support of this. Mayaud (1949), states that along the south coast of 
France between the Pyrenees and Provence intermediates occur between B. f 
fasciatus and B. cinereocapillus. Through the kindness of Dr. Mayaud we have 
examined four breeding males from the Etang de Salies and the Camargue, Mayaud's 
Nos. 729, 1059, 1066, and 1069. All these have a white stripe from base of bill to 
over and behind eye and are we consider B. f fasciatus. 

Budytes flavus dombrowskii Tschusi 

Budytes flavus dombrowskii Tschusi, 1903, Orn. Jb., 14: 161, Pantelimon, Rumania. 

Adult male : Differs from B. f. flavus in having the ear-coverts darker ; chin usually 
white ; upper throat often white. Wing 81-87, nm cl claw 8-10, tail 72-75 mm. Five 
males from the breeding area measured, a total of fifty-five specimens examined. 

Distribution : Breeding Rumania and Serbia in Yugoslavia ; in non-breeding season 
to Palestine, Iraq, and Africa as far south as the Sudan and Abyssinia. 

The female apparently differs from that of the nominate race in having rather 
darker ear-coverts, but we have seen no specimens from the breeding area and without 
these it is wellnigh impossible to give the comparative female characters. 

Budytes flavus plexus Thayer & Bangs 

Budytes flavus plexus Thayer & Bangs, 1914, Proc. New Engl. Zool. CI. 5: 41, Nijni Kolynsk, 
Kolyma, eastern Siberia. 

Adult male : A narrow white streak from base of bill to over and behind eye ; head, 
neck, and sides of face dark grey ; lores and ear-coverts blackish ; chin white ; throat 
yellow. Wing 83-84; hind claw n-12, tail 72-75 mm. Two males from breeding 



FROM WESTERN EUROPE TO WESTERN NORTH AMERICA 259 

area measured, a total of forty-four specimens examined. Thayer and Bangs give 
wing 81-82, tail 68-70 mm. for two males. 

The female has a duller grey head often with an olivaceous wash, and usually has 
the chest more or less washed with chrome yellow and some dark spotting. 

Distribution: Breeding northern areas of western and eastern Siberia as far west 
as the Petchora River ; in non-breeding season to Iraq, India, and China. 



Budytes flavus zaissanensis Poljakow 

Budytes flava zaissanensis Poljakow, 191 1, Messager orn. Mosk., 313: Lake Zaissan, west of 
Mongolian border, Turkestan. 

Adult male : A narrow white streak from base of bill to over and behind eye ; head 
slate grey; mantle olive-green; agreeing very closely with some specimens of B. 
thunbergi in these last two characters. Wing 77-84, hind claw 9-10, tail 67-78 mm. 
Two males from the breeding area measured, a total of nine specimens examined. 

We have not examined the female nor can we find any description of it. 

Distribution: Breeding Barnaul to junction of Altai and Irtysh Rivers and Lake 
Saissan; in non-breeding season to Sind, Punjab, and Bengal, India, Thailand, and 
West Java. 

Remarks: Poljakow compares this race to B. f. flavus and B. thunbergi. Sushkin, 
1925, gives wing 77-80, hind claw 9-3, tail 66-72-7 mm. 



Budytes flavus angarensis Sushkin 

Budytes flava angarensis Sushkin, 1925, Proc. Boston, Soc. Nat. Hist. 38: 33, Sharagolo-Kaia, 
Chikoi River, Transbaikalia. 

Adult male : A white streak from base of bill to over and behind eye ; mantle and 
borders of wing coverts less bright than B. f. zaissanensis. Wing 78-83, hind claw 
9-5-13, tail 72-74 mm. (Sushkin). 

Distribution : Breeding Lake Yevsi, Tunguzka and Angara Rivers to Lake Baikal, 
and the Chikoi River ; in non-breeding season to China and Thailand. 

We have not seen any specimens from the breeding area, but an adult male with 
a narrow white streak from base of bill to over and behind eye, which does not fit in 
with any other Eastern race, we consider is attributable to this race. It is from Pekin 
and was taken in May: Brit. Mus. Reg. No. 1949-9-243. Wing 80, hind claw 9, tail 
71 mm., and an adult male from Bangkok, Thailand, taken on 20 April 1931, now in 
Raffles Museum, agrees with this specimen. Sushkin has compared B. f. angarensis 
to B. flavus, B. simillimus, and B. zaissanensis. The two immature birds mentioned 
by Sushkin, 1925, taken in the Ordos area are probably of this race, but no date or 
measurements are given nor is any mention made about an eye stripe. The characters 
given are insufficient to determine them and both are evidently in immature dress as 
they are stated to be moulting from the young dress. It would therefore appear that 
they were bred in that area, and maybe B. f. angarensis breeds as far south as Ordos. 



260 ON THE SPECIES AND RACES OF THE YELLOW WAGTAILS 

Budytes flavus simillimus Hartert 

Budytes flava simillimus Hartert, 1905, Vog. Pal. Fauna, 1: 289, Kamschatka, Siberia. 

Adult male : A white streak from base of bill to over and behind eye ; head and 
nape rather darker grey than B. f. plexus ; lores and ear-coverts as in B. f. plexus ; 
mantle darker, more olive, less yellow-green. Wing 80-82, hind claw 9-5-11, tail 
64-69 mm. Three males from breeding area measured, a total of 319 specimens 
examined. The female has the head olivaceous ashy slightly contrasting with the 
mantle ; below, white or bufhsh-white ; lower belly washed with pale yellow. Hind 
claw often longer than the females of other races. The immature dress is ashy above, 
often slightly olivaceous ; rump greyer ; below, creamy white ; chest bumsh ; under 
tail-coverts often washed with pale yellow. In this dress it can be confused with 
Budytes citreola Pallas, though this species has a faint wash of yellow or buff on the 
forehead and the notch on the third primary 18-20 mm., up the feathers from the tip, 
this notch lying between the 7th and 8th primary, whereas in this race the notch on 
the 3rd primary is 15-16 mm., from the tip and lies between the 5th and 6th primary 
(see Sushkin, 1925). 

Distribution : Breeding Kamschatka ; in non-breeding season to India, as far west 
as the Punjab, Ceylon, Nicobar and Andaman Islands, China, Malay States, 
Philippine Islands, Dutch East Indies and New Guinea. 

Budytes flavus tschutschensis (Gmelin) 

Motacilla tschutschensis Gmelin, 1789, Syst. Nat. 2: 962, Tschutschi coast, Bering Strait, 
eastern Siberia, of which Budytes flavus alascensis Ridgway, 1903, Proc. Biol. Soc. Wash. 16: 
105, Western Alaska, is a synonym. 

Adult male : A white streak from base of bill to over and behind eye, broader than 
in B.f. plexus ; head and sides of face dark grey ; mantle darker than other races ; chin 
and upper throat usually white; dusky spots on lower neck; below, more lemon- 
yellow, not bright canary yellow as in B. f. simillimus and B.f. plexus. Wing 76-81, 
hind claw 10-11, tail 66-71 mm. Four males from breeding area measured, a total of 
twenty specimens examined. The female is similar to the male, but perhaps slightly 
duller. 

Distribution: Breeding north-eastern Siberia and Alaska; in non-breeding season 
to the Philippine Islands, West Java, and Dutch New Guinea. 

Remarks: The two males from the Philippine Islands and one male from Dutch 
New Guinea, Brit. Mus. Reg. No. 1888.7. 12.534 dated November, Brit. Mus. Reg. 
No. 1897. 12. 11.43 dated September, and Brit. Mus. Reg. No. 1916.5. 30.857, dated 
December, agree with this race in general colour and the paler, more lemon-yellow 
colour below and not with males of B. f. simillimus. 

Budytes luteus (Gmelin). Yellow Wagtail 

Budytes luteus luteus (Gmelin) 

Parus luteus Gmelin, 1774, (S. G.) Reise durch Russland, 3: 101, pi. 20, fig. 1, Astrakan, southern 
Russia, of which Motacilla campestris Pallas, 1776, Reise versch. Prov. Russ. Reichs, 3: 696 



FROM WESTERN EUROPE TO WESTERN NORTH AMERICA 261 

Russia; and Budytes flava var. flavifrons Sewertzow 1873, Vert. Geriz. Rashred. Turkest. 
Zhivoth, Mem. Soc. Amis Sci. Nat. Moscou, 8 (2) : 67. Turkestan (nom. nuda), 1875, Stray 
Feathers, 3: 424; and Budytes chlorocephalus Brehm, 1851, Naumannia, 2: 24, Reuthendorf, 
are synonyms. 

Adult male : Head yellow-green ; forehead, and streak from base of bill to over and 
behind eye, yellow. Wing 75-87, hind claw 8-1 1, tail 66-70 mm. Eight males from 
breeding area measured ; a total of 105 specimens examined. The female is difficult 
to distinguish from that of B. f. flavus, but it can be said that those with a more 
uniform head and mantle are this species and those with a greyish head contrasting 
with the mantle are B. f. flavus. 

Distribution: Breeding from the Volga River to the headwaters of the Yenisei 
River ; in non-breeding season to central, eastern, and southern Africa as far south as 
the Transvaal, Socotra Island, Arabia, India, and Ceylon. 

Budytes luteus taivanus Swinhoe 

Budytes taivana Swinhoe, 1863, Proc. zool. Soc. Lond.: 234, Formosa Island. 

Adult male : Top of head green, uniform with mantle ; lores to ear-coverts olivaceous 
black ; a broad yellow streak from base of bill to over and behind eye ; chin and throat 
bright yellow. Wing 76-87, hind claw 10-13, t an * 67-75 mm. The female differs from 
the male in being duller in colour. The immature dress can be distinguished from that 
of B. f. simillimus by the yellow in the eye stripe. Twenty males from breeding area 
measured, a total of eighty-two specimens examined. 

Distribution'. Breeding from the Lena River and Ija River west of Lake Baikal to 
the Amur River, also Sakhalin and Kurile Islands, in non-breeding season to Burma, 
China, Formosa, the Malay Peninsula, Borneo and Dutch East Indies. 

Budytes luteus flavissimus (Blyth) 

Motacilla flavissima Blyth, 1834, Loudon's Mag. 7: 342, England; of which Budytes rayi 
Bonaparte, 1838, Geog. <S- Comp. List Birds Europe & S. Amer. : 18, British Islands ; Budytes 
vema (S.D.W.) Wood, 1835, Analyst, 3: 31 ; 1836, 203 ; 1836,4 (16) : 296. Great Britain, nom. 
nuda; Budytes vema Wood, 1836, Brit. Birds: 219, Motacilla flaveola Temminck, 1835 Man. 
d'Orn. 2nd ed. 3 : 180, England, are synonyms. For other synonyms see Hartert, 1905, Vog. 
pal. Fauna, 1: 294. 

Adult male : Differs from B. I. luteus in having the forehead uniform in colour with 
the crown of the head. Wing 72-87, hind claw 8-1 1, tail 64-74 mm. The female is 
similar to that of B. I. luteus. Fifty-four males from breeding area measured, a total 
of 233 specimens examined. 

Distribution'. Breeding southern Norway, southern to eastern British Isles (rarely 
west Wales, Cornwall, and Devon), Heligoland in most years (Drost, 1948), western 
Holland, western Belgium, northern France, and Channel Islands ; in non-breeding 
season to Africa as far south as the Belgian Congo and Southern Rhodesia. 

Breeds alongside B.f. flavus in southern Norway (see Bernhoft-Osa, 1944 and 1946), 
and at Dunkirk, Pointe de Raguenes near Nevez, Finisterre, north-western France 
(see Mayaud, 1949, Ibis: 171). Dr. Holger Holgersen found it breeding in southern 
Norway in 1947 and 1949 and considers it to be a regular summer breeder. 

zoo. 1. 9. Mm 



262 ON THE SPECIES AND RACES OF THE YELLOW WAGTAILS 

Budytes superciliaris Brehm. White-browed Yellow Wagtail 

Budytes superciliaris Breton, 1854, /. Orn. Lpz.: 74, Khartoum, Sudan; of which Budytes 
leucostriatus Homeyer, 1878, 128, Lake Baikal area; Motacilla xanthophrys Sharpe, 1885, 
Cat. Birds. B.M. 10: 532, pi. 8, fig. 6, Lenkoran, Azerbaijan, southern Russia ; and Motacilla 
flava raddei Harms, 1909, Orn. Mber. 17: 2; Aschabad, Transcaspia, are synonyms. Hartert, 
i9°5» Vog. pal. Fauna, 1: 293, considers M. f. raddei to be an aberrant B. I. taivanus. 
Although we have not seen the type of Budytes leucostriatus, Homeyer gives the head as 
clear grey-black with a broad white stripe from the base of the bill to over and behind the 
eye. These characters agree with B. superciliaris and not with B. I. taivanus of which it is 
placed as a synonym by Hartert, 1905, Vog. pal. Fauna, 1: 298. 

Adult male: Top of head to nape jet black to grey-black ; nape often grey ; centre 
of crown to nape often olive-green ; a white or yellow streak from base of bill to over 
and behind eye; lores and ear-coverts black with usually some white flecking on 
latter and under eye ; chin white. Wing 77-85, hind claw 10, tail 66 mm. Two males 
from breeding area measured ; a total of thirty-five specimens examined. The female 
can be distinguished from that of other species by the grey head and mantle with only 
a slight wash of olivaceous green ; below, creamy white with a variable pale yellow 
wash from lower neck to under tail-coverts ; eye-streak buff or bumsh white ; some 
spotting at base of neck. The immature dress is very similar to that of the adult 
female. 

Distribution: Breeding southern Iran to Turkestan, also Bulgaria and eastern 
Yugoslavia; in non-breeding season to Egypt, the Sudan, Abyssinia, Arabia, and 
India. 

Col. Meinertzhagen (1949, Ibis: 472) records seeing a party of four males near Taif, 
Arabia, sometime between February and April. 

Budytes leucocephalus Przevalski. White-headed Yellow Wagtail 

Budytes leucocephalus Przevalski, 1887, Zap. Imp. Akad. Nauk. S.-Peterb. 55: 85, 1 Dzungaria, 
northern Turkestan. 

Adult male: Whole head to nape, chin and usually upper part of throat white, 
or white washed with grey ; sometimes a white eye-stripe is distinguishable. Wing 81, 
hind claw 10, tail 72 mm. One male from breeding area measured ; a total of thirty 
specimens examined. The female is similar to the male. The immature bird has the 
head and ear-coverts olivaceous grey ; a white streak from base of bill to over and 
behind eye; mantle washed with grey; below, chin and throat whitish washed 
with yellow ; a broken spotted collar at base of neck ; chest to under tail-coverts paler 
yellow than adult. 

Distribution : Breeding eastern Russia, Turkestan, and western Mongolia ; in non- 
breeding season to Africa as far south as north-eastern Northern Rhodesia and 
northern Nyasaland, Arabia and north-western India. 

Remarks : Sushkin found the character of the head constant in the breeding area 
at Lake Achit-Nor. The specimens we have examined have a remarkable close 
resemblance to each other and it should be noted that the sexes are alike, facts which 
have induced us to place it as a species and not as a very pale headed race of B. 
flavus. We have examined thirty-one adult specimens, four of which are females, 

1 This Russian version of the Memoires is apparently not available in Great Britain. 



FROM WESTERN EUROPE TO WESTERN NORTH AMERICA 263 

including one male in the Meinertzhagen Collection from Orox Nor in May, and 
compared them with the coloured Plate 10 of the male and female in Bianki 1905, 
Wiss. Res. Przevalski Cent. Asien, Z00L, 2, Vog. pt. 4, and find they agree very well 
with that plate, but it would appear the female figured is in immature dress. Finsch 
saw light-headed Yellow Wagtails, probably of this species, between the eastern end 
of Lake Zaissan and the Altai on 6 June, but none were obtained nor are they 
recorded as breeding there (see Suchkin, 1925). A description of this species was also 
published in Ibis, 1887: 401, but the one in the Russian journal has priority of date. 
Through the kindness of Dr. Barnard, Director of the South African Museum, we 
have had on loan the specimen recorded in /. 5. Afr. Orn. Un. 2: 92 (1906), from 
Kanyani, Northern Rhodesia, as 'Motacilla flava beema ? ' and find that it agrees 
perfectly with the Brit. Mus. series of this species: 

Budytes perconfusus Grant & Praed. Yellow-browed Yellow Wagtail 

Budytes perconfusus Grant & Praed, 1949, Bull. Brit. orn. CI. 69: 130, Khartoum, Sudan. 

Adult male: Above mantle rather darker than B. luteus and below rather paler 
yellow ; chin and throat yellow ; forehead to f orecrown grey ; a clear grey collar on 
hind neck ; crown olive-green ; a broad pale yellow streak from base of bill to over and 
behind eye ; lores to ear-coverts darker grey with white flecking. Differs from B. 
flava in the yellow streak over the eye and the paler grey head. Wing 78-85 ; hind 
claw 9-1 1 ; tail 68-70 mm. 

In fresh dress the forehead and nape is more washed with olivaceous green ; the 
mantle is darker ; the tips of the wing-coverts brighter yellow-green and only a few 
white flecks on the ear-coverts ; streak over eye yellow, not white or buff or washed 
with buff, as in B. flavus flavus in fresh dress. 

The female and young bird also have a yellowish eye streak. 

Distribution: Known only from scattered specimens from Frederikhavn, north- 
eastern Denmark, Pomerania, Germany, Wassenaar near The Hague, Holland, 
Abyssinia, the Sudan, and western Arabia. 

As stated above the five adult male specimens are exactly alike; the two from 
Khartoum, Brit. Mus. Reg. Nos. 1915. 12. 24.1429 and 1436 and Pomerania, Brit. 
Mus. Reg. No. 1941.5.30.819, were taken in April, the one from Denmark was taken 
on 3 May, and the one from Holland was taken in September. In the non-breeding 
season this species visits Abyssinia, the Sudan, and western Arabia and passes 
through Holland, Denmark, and Pomerania. A total of ten specimens examined, 
including a male from Fashoda, Sudan, March, Brit. Mus. Reg. No. 1902. 4. 20. 142; 
a male from Khartoum, Sudan, December, Brit. Mus. Reg. No. 1915. 12.24.1445 ; 
two males from Abyssinia, February and November, Brit. Mus. Reg. No. 1927. 11. 5. 
653, and 1934.8.9.352 ; and a female from Arabia, September, Brit. Mus. Reg. No. 
I 935-5- IO -78. We had considered placing it as a race of B. flava, but we feel that it is 
better treated as a species. We are confident that one day the breeding area will be 
discovered. 

B. No streak from base of bill to over and behind eye in male, though sometimes a 
short white mark behind eye : 
zoo. 1. 9. m m 2 



264 ON THE SPECIES AND RACES OF THE YELLOW WAGTAILS 

Budytes thunbergi (Billberg). Grey-headed Yellow Wagtail 
Budytes thunbergi thunbergi (Billberg) 

Motacilla thunbergi Billberg, 1828, Syn. Faun. Scand. 1 (2) Aves: 50, Lapland. For synonyms 
see Hartert, 1905, Vog. pal. Fauna 1: 291. 

Adult male : Head dark grey to near coal black ; no streak over eye, occasionally 
a short white mark behind eye ; more rarely a similar mark in front of the eye ; chin 
and throat yellow ; some spotting on lower neck in front. Wing 80-85, hind claw 8-1 1, 
tail 69-74 mm. Twenty-seven males from the breeding area measured ; a total of 
243 specimens examined. The female and immature are practically indistinguishable 
from that of B. flavus flavus, though perhaps some have rather a darker coloured 
top to the head. 

Distribution: Breeding northern Norway and northern Sweden to Finland and 
northern Russia as far east as the lower Yenisei River, also Estonia ; in non-breeding 
season to Africa as far south as Damaraland and the Transvaal, Arabia, India, 
Burma, and the Malay Peninsula. 

Remarks : S. Armington 1949, records at Ladugardsgarde, north-east of Stockholm, 
having observed a male Yellow Wagtail in the summer of 1947 which agreed perfectly 
with B. thunbergi. This bird was with a female which was indistinguishable from the 
female of B. f. flavus, but the nest was not located. About twenty pairs of B. f. flavus 
were breeding in the same locality. At the same place in 1949 Armington observed 
a male and female, the male agreeing with B. thunbergi, but had a superciliary streak 
over the right eye and a small patch behind the left eye. It is difficult to comment 
on the above, as there are no specimens to examine, but would remark that as 
B. thunbergi breeds in Finland and Estonia, it could be found breeding on the same 
latitude near Stockholm. 

As many females of both B. thunbergi and B. flavus are practically indistinguishable 
in skins and quite indistinguishable in the field, it cannot be said that the female 
with the Ladugardesgarde male was other than a B. thunbergi. As regards the male 
seen in 1949 as having a stripe over the right eye, this may have been a retention of 
the immature dress, but there is no proof that this is so. 

Jordans, 1923, mentions fifteen males of B. f. flavus taken near Upsala, Sweden, in 
May with varying coloured heads from pale grey to a darker or lighter crown, all 
having a superciliary stripe, and a series from Lapland which varies in a similar 
way, of which the lightest matches the darkest B.f. flavus, and others show all inter- 
gradations from this type to an almost black crown. Twenty per cent, have a light 
superciliary stripe indicated or even quite distinct. Count Gyldenstolpe has kindly 
picked out five representative specimens from this series and sent them to us for 
examination, and remarks in a letter to us dated 4 October 1949, ' B. f. thunbergi is 
found at Upsala during its migrations in May, together with B. f. flavus, hence the 
statements made by Jordans'. 

All these five specimens are clearly B. thunbergi thunbergi; the Lapland ones all 
taken in June and the Upsala ones on 6 and 12 May. One from Lapland, taken on 
16 June 1 93 1, which has the dark head and ear-coverts of B. t. thunbergi has an 
indication of a white streak before as well as behind eye, but not over the eye. All 



FROM WESTERN EUROPE TO WESTERN NORTH AMERICA 265 

have some spotting on the lower neck in front. It would appear that Jordans con- 
sidered this Lapland specimen of 16 June 1931 as an intermediate between B. flavus 
and B. thunbergi, but we are satisfied that this specimen is B. thunbergi, and not an 
intermediate between that species and B. flavus. 

Budytes thunbergi cinereocapillus (Savi) 

Motacilla cinereocapilla Savi, 1831, Nuovo Giorn. Lett. Pisa, 22, 190, Tuscany, Italy. 

Adult male : Differs from the nominate race in having the chin and neck in front 
white, or rarely some yellow mixed with the white on the lower neck in front. Wing 
81-83, nm d c l aw 9-1 1, tail 70-72 mm. Nine males from breeding area maesured, a 
total of thirty-one specimens examined. The female and immature are practically 
indistinguishable from those of B. f. fasciatus, both having a streak from the base of 
the bill to over and behind eye. 

Distribution: Breeding southern France to Italy, Dalmatia, Switzerland, and 
Algeria ; in non-breeding season to western, northern, and eastern Africa as far south 
as Senegal and Uganda, also Arabia. 

Kirkman & Jourdain, 1913, British Bird Book, Vol. 4: 477, state that this race 
breeds in Tunis, and this has been quoted by other authors. We cannot find any 
evidence in support of this. 

Remarks'. Thonen (1948) records and figures the head of a Yellow Wagtail breeding 
at Lake Neuenburger, near Basle, Switzerland. 

This figure and the description agrees well with specimens in the British Museum 
collection of Budytes thunbergi cinereocapillus which has in some specimens the pure 
white more confined to the throat and a white fleck behind the eye. This figure was 
drawn from life, as the bird was not collected. Ticehurst and Whistler (1927) state 
that this race is found in the plains and B. f. fasciatus in the mountains, but Tice- 
hurst obtained a male of B. f. fasciatus in June at Argeles-sur-Mer. It would thus 
appear that both breed in the same area. The single record of this race from Great 
Britain can be accepted. There is little doubt that it was collected near Marazion 
Station. The specimen has disappeared, but there is an excellent coloured figure of 
it in Gould's Birds of Gt. Britain, 3: pi. 5, 1873. 

Budytes thunbergi pygmaeus Brehm. 

Budytes pygmaeus Brehm, 1854, /. Orn. Lpz.\ 74, (note), north-east Africa. 

Adult male : Similar to B. t. cinereocapillus but smaller in size ; chin and throat 
sometimes yellow ; often an olive-green patch on crown of head and a white streak 
behind eye. Wing 70-78, hind claw 8-10, tail 58-67 mm. The female is very similar 
to that of B. t. cinereocapillus, but is smaller ; sometimes a white mark behind eye. 
Thirty-six specimens examined. 

Distribution: Egypt. 



Budytes thunbergi macronyx Stresemann. 

Budytes flavus macronyx Stresemann, 1920, 
Russia. 

Adult male : Differs from the nominate race in having a rather darker mantle and 



Budytes flavus macronyx Stresemann, 1920, Avifauna Macedonica: 76, Vladivostock, eastern 
Russia. 



266 ON THE SPECIES AND RACES OF THE YELLOW WAGTAILS 

a longer hind claw ; chin white ; sometimes a short white mark behind eye. Wing 
78-84, hind claw 10-15, tail 69-75 mm. Fourteen males from breeding area measured, 
a total of seventy-one specimens examined. The female has the head olivaceous green 
or olivaceous grey ; sometimes a short light mark behind eye. The immature dress is 
grey or olivaceous grey above and creamy white below. 

Distribution: Breeding Siberia; in non-breeding season in China, Siam, Burma, 
Indo-China, Philippine Islands, Singapore, Borneo, Sumatra and Java. 

Budytes thunbergi alakulensis Grant & Praed. 

Budytes thunbergi alakulensis Grant & Praed, 1949, Bull. Brit. orn. CI., 69: 131, Lake Alakul, 
Turkestan. 

Adult male : Similar to Budytes thunbergi thunbergi, but head rather darker, more 
coal black. Wing 80, hind claw 8, tail 68 mm. 

Distribution : Breeding Lake Alakul ; in non-breeding season to Kiukiang, Yangtse 
River, China. 

Two specimens examined. 



Budytes feldegg (Michahelles) . Black-headed Yellow Wagtail 

Motacilla feldegg Michahelles, 1830, I sis: 812, Split, Dalmatia, Yugoslavia; of which Budytes 
melanogrisea Homeyer, 1878, /. Orn. Lpz.\ 128, India; Budytes aralensis Homeyer, 1878, 
J. Orn. Lpz. : 128, Lake Aral (compared to B. feldegg, head given as coal black ; below lemon 
yellow); Budytes flava suschkini Domaneiwski, 1925, Ann. Mus. Zool. Polon, 4: 95 & 107, 
no type locality; and Motacilla kaleniczenkii Kaleniczenko, 1839, Bull. Soc. Nat. Moscow. 
229, pi. 20, Crimea, are synonyms. 

Adult male: Forehead to nape, sides of face, ear-coverts and sides of neck jet black 
to coal black ; sometimes variable white streak between black face and yellow throat ; 
chin and throat yellow; often some green on top of head and some grey at nape. 
Can be distinguished from B. superciliaris in having no streak from base of bill to 
over and behind eye. Wing 78-85, hind claw 9-1 1, tail 66-76 mm. Thirty-three 
males from breeding area measured ; a total of 377 specimens examined. The female 
and immature plumages are similar to those of B. superciliaris, but there is no streak 
from the base of the bill to over the eye, though sometimes there is a short light mark 
behind the eye. 

Distribution: Breeding in Montenegro, Serbia in Yugoslavia, Albania, Greece, 
Turkey, and Syria to the Black and Caspian Seas, Lake Aral, north-western Iran, and 
Turkestan ; in non-breeding season to southern France, eastern Africa as far south as 
Uganda and Kenya Colony, southern Arabia, Socotra Island and India. 

In adult females there is considerable individual variation on the head which does 
not appear to have any relation to the breeding and non-breeding season. These 
variations are well shown in PI. 1, Ibis, 1932, though the black colouring" in figs. 3, 
4, and 5 is much too dull. Rarely males have the mantle grey with a slight olivaceous 
wash ; below, chin and throat white with a faint touch of yellow ; rest of underparts 
very pale yellow. A plumage very similar to some adult females. The four specimens 



FROM WESTERN EUROPE TO WESTERN NORTH AMERICA 267 

recorded from Great Britain have disappeared. We are of opinion that the claim 
that they were British taken should be viewed with grave suspicion. 

We have to thank Dr. K. H. Voous for much kind help, and Dr. Holger Holgersen 
for translations from Norwegian journals. 

REFERENCES 

Alexander, H. G. 1950. Variant- Yellow Wagtails, Brit. Birds 43: 31. 

Armington, S. 1949. Motacilla flava thunbergii Billb. : Stockholm, Dansk. Orn. Foren. Tidsskr. 

43: 92. 
Baker, E. C. S. 1926. Fauna of British India, Birds, 2nd ed. 3: 267, London. 
Bernhoft-Osa, A. A. 1945. Hornugle — Asio otus — og andre sjeldnere fugler pa Jaeren, Stavanger 

Mus. Arsh. 1944: 138-140. 

1946. Engelsk Gulerle, en ny Rugefugl for Norge, Naturen: 317. 

Degland, C. D. & Gerbe, Z. 1867. Ornithologie europeenne, 2nd ed. 1: 380. 
Dementiev, G. P. 1934. Systema Avium Rossicarum, Oiseau, Paris 4: 606. 

1937. Polnuii opredelitel Ptitz S.S.S.R. 4: 142, Moskva & Leninghrad. 

Domaniewski, J. 1925. Systematik und geographische Verbreitung der Gattung Budytes Cuv. 

Ann. Mus. zool. polon. 4: 85. 
Geroudet, P. 1946. Le passage des Bergeronnettes printanieres du Nord de l'Europe, Nos 

Oiseaux: 202. 
1948. Sur l'apparition de trois sous-especes de la Bergeronnette printaniere en Suisse, Nos 

Oiseaux: 284. 
Glegg, W. E. 1931. The birds of L'lle de la Camargue et la Petite Camargue, Ibis: 220. 
Grant, C. H. B. & Mackworth-Praed, C. W. 1939. On the races of Motacilla flava occurring in 

Eastern Africa, Bull. Brit. orn. CI. 59: 160. 
1942. On the conspeciflc status of Budytes flava, B. luteus & B.feldegg, Bull. Brit. orn. 

CI. 62: 58. 
Grote, H. 1919. Ornithologische Beobachtungen aus dem siidlichen Uralgebiete, /. Orn. Lpz. 

67: 371. 

1937. t)ber Motacilla flava mutatio lutea, Orn. Mber. 163. 

Harrison, J. G. 1945. Some remarks on the problem of Sykes' Wagtail in the British Isles, 

Ibis: 69. 
Hartert, E. 1922. Die Vogel der palaarktischen Fauna 3: 2097, Berlin. 

1932. Die Vogel der palaarktischen Fauna . . . Ergdnzungsband: 146, Berlin. 

Homeyer, E. F. v. 1878. Beitrage zur Gattung Budytes, J. Orn. Lpz.: 126. 

Ivanov, A. J. 1935. tiber die Formen der Gattung Budytes, C.R. Acad. Sci. U.R.S.S. 3: 277. 

Johansen, H. 1946. De Gule Vipstjerters (Motacilla flava) Systematik og Udbredelse, Dansk. 

Orn. Foren. Tidsskr. 40: 121. 

1949. En aberrant Gul Vipstjert (Motacilla flava), Dansk. Orn. Foren. Tidsskr. 43: 93- 

Jordans, A. v. 1923. Tiber seltenere und iiber fragliche Vogelformen meiner Sammlung, Falco, 

19: 15 (Sonderheft). 
Jourdain, F. C. R. 1915. Notes on Bird Life of Eastern Algeria, Ibis: 142. 
Kleiner, A. 1935. Die Rassen der Schafstelzen in Ungarn, Budapest. 
1936. Mitteilungen iiber die Schafstelzen (Motacilla, Aves) Bulgariens und seiner angren- 

zenden Gebiete, Mitt, naturw. Inst. Sofia 9: 69. 
1939. Des races de la Bergeronnette (Motacilla flava) au Bassin des Carpathes, Festschrift 

f. Embrik Strand 5: 365. 
Lynes, H. 1925. Contributions to the Natural History of Morocco, Mem. Soc. Sci. Nat. Maroc. 

13, pt. I: 43 . 
Mayaud, N. 1936. Inventaire des Oiseaux de France: 140-141, Paris. 
1949. The races of Motacilla flava, Ibis: 171. 



268 ON THE SPECIES AND RACES OF THE YELLOW WAGTAILS 

Poljakow, G. J. 191 1. Eine neue Form der Schafstelze, Messager Orn., Mosk.: 313. 

Riley, J. H. 1918. Annotated Catalogue of a Collection of Birds made ... in NE. Siberia, Proc. 

U.S. Nat. Mus. 54: 621. 
Smith, S. 1950. The Yellow Wagtail, London. 

Sowerby, A. de C. 1 923. The Naturalist in Manchuria 3: 177, Tientsin. 
Stresemann, E. 1920. Avifauna Macedonica: 74, Miinchen. 
Sushkin, P. P. 1914. Die Vogel der Mittleren Kirgisensteppe, /. Orn. Lpz. 62: 329. 

1924. Exhibition of eggs of Budytes flava leucocephala, Bull. Brit. orn. CI. 45: 39. 

1925. Notes on systematics and distribution of certain Palaearctic Birds. Proc. Boston 

Soc. Nat. Hist. 38: 33. 
Thayer, J. E. & Bangs, O. 191 4. Notes on the birds and mammals of the Arctic coast of East 

Siberia, Proc. New Engl. Zool. CI. 5: 41. 
Thonen, W. 1948. Eine Schafstelzenbrut am Famel (Neuenburgersee), Orn. Beob. 45: 38. 
Ticehurst, C. B. 1922. Notes on Indian Wagtails, /. Bombay Nat. Hist. Soc. 28: 1082. 

& Whistler, H. 1927. On the summer Avifauna of the Pyrenees Orientales Ibis: 294. 

1932. On the ornithology of Albania, Ibis: 53. 

Ticehurst, N. F. 1907. On the Yellow Wagtails and their position in the British Avifauna, 

Brit. Birds 1: 133. 
Voous, K. 1950. The races of Yellow Wagtail wintering in the Indo- Australian Archipelago, 

Treubia 20: 647. 
Whistler, H. 1940. The White-headed Wagtail (Motacilla flava leucocephala), Ibis: 335. 



PLATE 34 PLATE 35 

j. Budytes flavus flavus 12. Budytes luteus luteus 

2. Budytes flavus beema 13. Budytes luteus taivanus 

3. Budytes flavus fasciatus 14. Budytes luteus flavissimus 

4. Budytes flavus dombrowskii 15. Budytes perconfusus 

5. Budytes flavus plexus 16. Budytes thunbergi thunbergi 

6. Budytes flavus zaissanensis 17. Budytes thunbergi cinereocapillus 

7. Budytes flavus angarensis 18. Budytes thunbergi pygmaeus 

8. Budytes flavus simillimus ig. Budytes thunbergi macronyx 

9. Budytes leucocephalus 20. Budytes thunbergi alakulensis 

10. Budytes flavus tschutschensis 21. Budytes feldegg 

11. Budytes superciliaris 22. Known breeding distribution of all the Yellow Wagtails 



Bull. B.M. {N.H.) Zoology, 1, 9 



PLATE 34 








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PRINTED IN 

GREAT BRITAIN 

AT THE 

UNIVERSITY PRESS 

OXFORD 

BY 

CHARLES BATEY 

PRINTER 

TO THE 

UNIVERSITY 



V. 



1 8 DEC 1952 

MAMMALS COLLECTED BY 

MR. SHAW MAYER 

IN NEW GUINEA 
1932-1949 



ELEANOR M. O. LAURIE 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ZOOLOGY Vol. 1 No. 10 

LONDON: 1952 



MAMMALS COLLECTED BY 

MR. SHAW MAYER 

IN NEW GUINEA 
1932-1949 

BY 

ELEANOR M. O. LAURIE 



... M6 




Pp. 269-318 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ZOOLOGY Vol.i No. 10 

LONDON : 1952 



THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted in ig4g, is issued 
in five series, corresponding to the Departments of the 
Museum. 

Parts appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

This paper is Vol. i, No. 10 of the Zoological series. 



PRINTED BY ORDER OF THE TRUSTEES OF 
THE BRITISH MUSEUM 

Issued December 1952 Price Fifteen shillings 



MAMMALS COLLECTED BY MR. SHAW MAYER 
IN NEW GUINEA, 1932-1949 

By ELEANOR M. O. LAURIE 

SYNOPSIS 

This paper gives a detailed account of a large collection of Mammals, mainly Marsupials and Rodents, 
from north-east New Guinea and eastern Papua (south-east New Guinea). Comparative descriptions 
are made of 13 new forms comprising 1 new genus (rodent), 7 new species (3 marsupials, 2 rodents, 1 bat, 
and 1 monotreme), and 5 subspecies (3 marsupials and 2 rodents). 

During the years 1932-1949 Mr. Shaw Mayer made a collection of mammals in 
New Guinea. Most of the specimens came from localities of comparatively high alti- 
tude, where the hill-sides are covered with rain forest. Between 5,000 and 8,000 ft. 
the lower limit of the wet mossy forest is often reached. From 10,000 to 11,000 ft. 
is a drier zone of grassland and coniferous forest, the upper limit of the forests being 
at about 14,000 ft. A list of all the localities from which specimens were obtained 
is given in Appendix II. Most of them are in north-east New Guinea: the Hagen 
Range and Sepik-Wahgi Divide, 4,500-8,500 ft. ; the Kratke Mountains and Upper 
Waria River district, 2,500-6,000 ft. ; the Upper Ramu River Plateau, 6,000 ft. ; 
Mount Wilhelm and Herowagi, Bismarck Range, 6,000-10,000 ft. ; the Ramu 
Purari Divide which is south-east of the Bismarck Range, 7,500-8,000 ft. ; and in 
eastern Papua, south-east New Guinea: Mount Simpson, Mount Mura (30 miles 
NW. of Mt. Simpson) and the Maneao Range (35 miles NW. of Mt. Simpson), 1,000- 
7,000 ft. (see Fig. 1). A few specimens, mainly rodents, were also collected from 
West Fergusson Island (which is about 40 miles from the mainland), between 600 
and 3,000 ft. Many of these regions have not been investigated before, particularly 
those near the Bismarck Range and Mount Simpson. 

The collection comprises 370 marsupials belonging to 29 species, 380 rodents 
belonging to 31 species, 31 bats belonging to n species, and 5 monotremes belonging 
to 3 species. 

Among these specimens which are dealt with in this paper are 13 new forms: 
6 marsupials (3 species and 3 subspecies), 5 rodents (1 genus, 2 species, and 2 sub- 
species), 1 bat (species), and 1 monotreme (species). 

Most of the recent work on mammals of New Guinea has been done by G. H. H. 
Tate (1935-1951), using several valuable collections from Vogelkop, the Arfak 
Mountains, Humboldt Bay, the Weyland Mountains, Mount Wilhelmina, the Inden- 
burg River, Fly River, Oriomo River, the Central Division of Papua, the Huon 
Peninsula, and from islands off the coast of New Guinea. An account of the rodents 
of Australia and New Guinea by Tate (1951) has been published while this manuscript 
was in the press. I have, however, been able to refer to it and in the main have fol- 
lowed his revised nomenclature. Accounts of many of the forms from New Guinea 
have been given by Thomas and also by Schlegel, Milne-Edwards, Matschie, Ram- 
say, Rothschild & Dollman, Hinton & Ellerman. 



272 



MAMMALS COLLECTED BY MR. SHAW MAYER 



This account includes references to a few co-types and paratypes which have 
already been mentioned in their type descriptions but are included here as they are 
part of this collection. 



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Fig. 1. Map of eastern New Guinea showing localities near which specimens were obtained. 

Amongst the commonest animals collected are the following : 
Marsupials: Eudromicia caudata, the Long-tailed Dormouse Phalanger; Dacty- 
lopsila trivirgata melampus, the Black-footed Striped Phalanger ; Dactylonax palpator, 
the Long-fingered Striped Phalanger; Phalanger vestitus, a Cuscus; Pseudocheirus c. 
cupreus and P. c. corinnae, Ring-tailed Opossums (only from NE. New Guinea but 
previously recorded from both NE. and SE.) ; Peroryctes longicauda omata, the 
Ornate Bandicoot (only from NE. New Guinea but previously recorded from both 
NE. and SE.) ; and Satanellus albopunctatus , the northern Native Cat which is now 
regarded as being synonymous with the southern form daemonellus. The Ring-tail 
Opossum Pseudocheirus forbesi larvatus also appears to be fairly common but is 
restricted to north-east New Guinea. 



IN NEW GUINEA, 1932-1949 273 

Rodents: Rattus exulans browni, Brown's Island Rat, a small rat common in 
native huts ; Rattus ruber tramitius, a common outdoor scavenger in native gardens 
and sometimes in huts ; Melomys rufescens rufescens, a Mosaic-tailed Rat ; Pogonomys 
mollipilosus, Pogonomys sylvestris, and Pogonomys macrourus, Prehensile-tailed 
Rats; and Mallomys rothschildi, a giant rat. Melomys fellow si, a Mosaic-tailed Rat, 
Crossomys moncktoni, Monckton's Water-rat, Hyomys goliath goliath, one of the 
giant rats, and Parahydromys asper, a water-rat are also common, but in this 
collection have only been taken from north-east New Guinea though their range, 
apart from Melomys fellowsi, extends into north-east Papua. 

Only a small number of bats were collected. They include, however, one new 
species of Otomops, which is of interest as this is only the second time that the genus 
has been recorded from New Guinea. 

The Echidnas include one new species of Zaglossus. 

The fauna of New Guinea is closely related to that of Australia. The great majority 
of the forms, however, are specific to New Guinea and the neighbouring islands. 

Throughout this paper the specimen numbers given are the British Museum 
registered numbers unless otherwise stated. 

Where there are a large number of specimens of a species the extremes, mean, 
and standard deviation which gives some idea of the variation from the mean, are 
given instead of the detailed measurements of each specimen. 

I should like to take this opportunity of expressing my thanks to my colleagues 
in the Mammal Room for their help, especially to Dr. T. C. S. Morrison-Scott and to 
Mr. R. W. Hayman, who helped with the identification of the bats and has described 
the new Otomops in this paper. 

MONOTREMATA 

Zaglossus bartoni bartoni (Thomas) 

Acanthoglossus bruijnii bartoni Thomas, 1907, Ann. Mag. Nat. Hist. 7: 294. 
Type locality: Mount Victoria, Papua, 8,000 ft. 

Two specimens, ?$ 50.1453, $ 1452, from Bubu River district, NE. New Guinea. 

Measurements in mm. (taken in the flesh) [Female first, male second]: Total 
length 600, 573; hind foot — , 62; weight 2i| lb., 13 lb.; length of skull 175, 169; 
basal length 167, 158; breadth of braincase 55, 57; muzzle from level of lacrymal 
canal 110-3, 106*1 ; gnathion to back of palatal bones 154, 146 ; least inter-orbital 
breadth 20-0, 18-2; width of rostrum 40 mm. from tip n-8, 11-4. 

Zaglossus bubuensis sp. n. 

Type locality: Bubu River district, NE. New Guinea, c. 7,000-8,000 ft. 

Type: Adult £ 50.1454, collector's No. 544, 8 Nov. 1936. Skin and skull. 

Similar to bartoni in having five claws on all the feet, in the spineless undersurface 
which, however, is only thinly covered with hair, and in the uniform whiteness of 
the short (max. length c. 32 mm.) spines. It differs from Z. b. bartoni in that its hair 
is brown, not black, and does not quite cover the spines on its back. The hair on the 
backs of all four feet is light brown. 



274 



MAMMALS COLLECTED BY MR. SHAW MAYER 



Body measurements in mm. (taken in the flesh) : Total length 656 ; hind foot 60 ; 
weight 17P). 

The size and shape of the skull is somewhat similar to that of Z. b. bartoni, but 
the rostrum is not so curved. 

Skull measurements in mm. : 











"§ 






g 






.3 


8 


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1 S 




5 




§ 8 


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Type of bubuensis 


177 


167 


57-5 


108 


156 


17-1 


12-4 


Type of bartoni 


184 


174 


59'5 


115 


161 


20-0 


130 



Tachyglossus aculeata lawesi (Ramsay) 

Echidna (Tachyglossus) lawesi Ramsay, 1877, Proc. Linn. Soc. N.S.W., 2: 32. 
Type locality: Port Moresby, SE. New Guinea. 

Juvenile $50.1450 (skull and piece of skin), Apimuri, Kratke Mts., NE. New 
Guinea; skull $ 50.1451, locality unknown but probably same as 50.1450. 

MARSUPIALIA 

Thylogale bruijni browni (Ramsay) 

Halmaturus brownii Ramsay, 1877, Proc. Linn. Soc. N.S.W. 1: 307. 

Type locality: New Ireland. 
Macropus lugens Alston, 1877, Proc. Zool. Soc. Lond. 1877: 126. 

Type locality: Duke of York Island or adjoining shores of New Britain or New Ireland. 
Macropus tibol Miklouho-Maclay, 1885, Proc. Linn. Soc. N.S.W. 10: 141. 

Type locality: 'Maclay Coast' north of Finisterre Range and east of Madang. 
Thylogale lauterbachi Matschie, 1916, Mitt. Zool. Mus. Berl. 8: 290-292. 

Type locality : Ogeramnag near Source of Bulung River. 
Thylogale brunii brownii Ramsay, Tate, 1948, Bull Amer. Mus. Nat. Hist. 91: 319-320. 

Five specimens from NE. New Guinea: <$ 50.1445, Arau, Kratke Mts. ; juv. <J 1447, 
$1446, Buntibasa district, Krakte Mts.; ^1449 (skull and piece of skin), Kam- 
baidam, Kratke Mts. ; $ 1448, Bubu River district. 



Dorcopsulus vanheurni vanheurni (Thomas) 

Dorcopsis vanheurni Thomas, 1922, Ann. Mag. Nat. Hist. 9: 264. 

Type locality: Doormanpad-bivak, N. New Guinea, 1410 m. 
Dorcopsulus vanheurni vanheurni Thomas, Tate, 1948, Bull. Amer. Mus. Nat. Hist. 91: 286—287. 

Six specimens. Three from NE. New Guinea: S 50-*434> Kuraka, Kratke Mts.; 
$ 1443, $ 1444, Saiko, Bubu River ; and three ?$ii4o, 1141, juv. 1142, from Boneno, 
Mt. Mura, eastern Papua. 

Two females, Nos. 1140 and 1141, and a male, No. 1434, are of interest as their 
pelage appears to be that of the fully adult animal, which differs from that of the 



IN NEW GUINEA, 1932-1949 275 

type, a young adult female, in being shorter, thinner, and of darker grizzled brown 
without the somewhat rufous colour. 

Measurements in mm. (taken in the flesh) : 





















"»» 












?s 














« 












•^3 












-ss 






















-*s 




5ya 


53 












t3 






^ 
^ 




g 5 


^s 


•2* 




"a 








53 




s 


£» 




5^o <S 




13 « 








fe 


m 


£ 


S3 


53 


53 

05 


N .G 


53 




•fc, 


§ 


^ 


50.1140 


$ 


446 


347 


107 


40 


74-9 


46*I 


33'3 


4-0 


27-8 


13-6 


9-6x3-5 


1141 


¥ 


413 


320 


103 


40 


7°'5 


4 2-7 


32-1 


4-6 


28-0 


13-6 


9-6x3-4 


1434 


0* 


341 


402 


IOO 


39 


74-0 


44-2 


33-o 


5'4 


27*0 


I3'5 


9-0x3-5 


1443 


s 


395 


298 


96 


37 


69-1 


43-2 


3i-8 


2-5 


26-3 


I3'4 


8-i X3-2 


1444 


V 


375 


295 


98 


3« 


70-0 


42-7 


3i-5 


4-6 


26-9 


13-4 


8-5X3-5 



Dendrolagus dorianus dorianus Ramsay* 

Dendrolagus dorianus Ramsay, 1883, Proc. Linn. Soc. N.S.W. 8: 17. 
Type locality : Mount Astrolabe, SE. New Guinea. 

Dendrolagus dorianus dorianus Ramsay, Rothschild & Dollman, 1936, Trans. Zool. Soc. Lond. 
21: 477-549- 

Nine specimens. Seven from NE. New Guinea: ^50.1427, 1428, 1423, juv. 
<J 1422, ? 1424, 1426, juv. $ 1425, south side Bubu River, NE. New Guinea; and 
two from eastern Papua: $ 1143, Enaena, NE. slopes Mt. Simpson; and $ 1144, 
Manaeo Range. 

The colour of the pelage of these specimens varies from a dark brown, e.g. No. 
1428, to a light greyish -brown, No. 11.43. 

Although the tail is non-prehensile the hairs on it in specimens Nos. 1143 and 1144 
are worn down so that they are quite short and bristly. The tails of one or two of the 
specimens of Dendrolagus already in the Museum's collection also have this appearance. 

Measurements in mm. (taken in the flesh) : 



K 




1 








so 

1 


.0 
















I 


00 


1^ 


1 


3 


1 




N .0 


1 


"s 


*, 


"s 


§ 


"s 


S 


50.1143 


? 


596 


662 


117 


48 


115-8 


72-0 


47-5 X 22-8 


13-3 


10-9 x 6-8 


6-5 x 6-6 


6-8x6-7 


7-ox 7-2 


7-1x7-0 


1426 


? 


600 


490 


102 


50 


108-8 


65-8 


46-5 X 22-8 


13-2 


IO-2X 6-2 


6-3x6-4 


6-7x6-6 


6-9 x 6-9 


7-1 x 6-2 


1424 


$ 


615 


463 


IOO 


50 


106-9 


67-2 


44-0 x 22-4 


12-5 


9-8x5-8 


6-0x5-9 


6-5 x 6-3 


6-7 x 6-5 


6-4 x 5-8 


1423 


6" 


628 


497 


I07 


57 


113-2 


75-5 


46-9X24-5 


13-5 


10-4 x 6-5 


6-5x6-4 


7-0x6-8 


7-5x6-8 


7-5x6-7 


1 144 





633 


550 


IO3 


45 


109-0 


68-8 


43-6x23-5 


13-6 


10-4 x 6-7 


6-6x6-3 


6-9 x 6-7 


7-1x6-9 


7-IX6-5 


1427 


5 


687 


586 


115 


53 


124-0 


77-o 


51-7x26-8 


14-0 


11-0x7-2 


6-7x6-4 


7-3X6-5 


7-2x6-8 


7-1x6-5 


1428 


a 


730 


570 


117 


57 


122-7 


77-5 


49-0x24-9 


13-6 


10-3 x 6-6 


6-6x6-1 


7-0x6-5 


7-5x6-8 


7-5X6-7 



Dendrolagus dorianus shawmayeri Rothschild & Dollman 

Dendrolagus goodfellowi shawmayeri Rothschild & Dollman, 1936, Trans. Zool. Soc. Lond. 21: 
484, 486. 
Type locality: Kratke Mts., NE. New Guinea, 4,500 ft. 
Dendrolagus dorianus shawmayeri Rothschild & Dollman, Tate, 1948, Bull. Amer. Mus. Nat. 
Hist. 91: 237-351. 

Six specimens. Five from NE. New Guinea: two co-types, juv. $ 50.1429, $ 1430 
(skull only), from Arau, Kratke Mts.; juv. $ 1431 (skin only), Binemarian, Kratke 

* For D. d. notatus, see Appendix III, p. 318. 



276 MAMMALS COLLECTED BY MR. SHAW MAYER 

Mts. ; sex 50.1432 (flat skin only), north side Bubu River; $ 1814, near Herowagi, 
south slopes Bismarck Range ; and one flat skin (no number) from mountains of SE. 
New Guinea, behind the island of Samaria. 

No. 1429 is the co-type mentioned by Rothschild & Dollman (1936), who also 
refer to another specimen, presumably No. 1431. 

Measurements in mm. (taken in the flesh) : 



I 


to 


1 
« 

If 


§ 




1 


1 


1 


11 
$1 


1 


1 


*, 




1 


°g 


5 


50.1429 
1430 
1814 


C?* 

$ 


480 


675 


"3 


51 


92-0 
99-5 
100-3 


59-7 
61-3 

6 3 -7 


44-9x15-3 
46-1 x 21-3 

44-OXI9-7 


II-2 
II-O 

n-9 


9-8x5-5 
9*9x4-8 
9-6x5-0 


5-4X4-9 
5-3X5-I 
5-7X5-3 


5-8x5-1 
5-7x5-2 
6-0x5-5 


6-1x5-0 
6-1x5-2 
6-2 x 5-5 


6-ox 5-0 
6-1x5-2 
6-0x5-4 



* juvenile 

Distoechurus pennatus neuhaussi Matschie 

Distoechurus neuhaussi Matschie, 1916, Mitt. Zool. Mus. Berl. 8: 292. 

Type locality: Sattelberg Mts., Huon Gulf, Dutch New Guinea. 
Distoechurus pennatus amoenus Thomas, 1920, Ann. Mag. Nat. Hist. 6: 537. 

Type locality: Rawlinson Mts., New Guinea. 
Distoechurus pennatus neuhaussi Matschie, Tate & Archbold, 1937, Bull. Amer. Mus. Nat. 
Hist. 73: 388-390. 

Ten specimens. Four from eastern Papau: $ 50.1073, $ 1076, 1074, 1075, Enaena, 
(1076 from Ikara), NE. slopes Mt. Simpson ; and six from NE. New Guinea: $ 1387, 
$ 1388, 1389, Buntibasa district, Kratke Mts. ; and $ 181 1, juv. $ 1812 (in pouch 
of 1811), Yandara, Bismark Range; $ 1813, Guyebi, Bismarck Range. 

This series extends the range of neuhaussi from the Sepik River area, NE. New 
Guinea (Tate & Archbold, 1937) to Mt. Simpson, eastern Papua. Very slight differ- 
ences in the general colour of the specimens from the three localities can be noted. 
Those from eastern Papua are a uniform light brown, those from the Kratke Mts. 
slightly darker and more ochraceous, and those from Yandara and Guyebi (Mt. 
Wilhelm) slightly darker and greyer, though No. 1813 is hardly distinguishable 
from those from eastern Papua. 

Measurements in mm. (taken in the flesh) : 

























•~<> 






^ 


















« 






■5 
















-« 








►0 








^e 




«s 




§> 








s 

^3 









8 


<o 


rO 






N 








< 

^ 




"<§ 




^ 




4§ 


S 

1 I 


00 


«5 




§ 


s< 


<o 


<ao * 


•S Q 






t2 <S 


s 
fc 




s 


5 






N -o 






8 


s £ 

^ ^ 


50.1075 


? 


108 


137 


20 


12 


26-1 


17-2 


5-8 


11-5x3-8 


15-4 


3-2 


1076 


¥ 


109 


136 


20 


11 


25-9 


17-2 


5-8 


11-4x3-3 


16-0 


3'2 


1074 


¥ 


no 


152 


21 


n 


27-0 


17-1 


57 


11-6x3-9 


16-0 


3-3 


1073 


6* 


113 


145 


21 


12 


26-8 


17-7 


6-o 


12-8x3-5 


16-5 


3-3 


1387 


0* 


103 


136 


20-5 


12 


26-9 


I7'5 


6-o 


— X3-4 


— 


3-5 


1389 


¥ 


109 


149 


21 


12 


27-9 


19-2 


67 


12-7x3-7 


16-5 


3-6 


1388 


? 


116 


142 


21 


13 


28-6 


19-2 


6-5 


12-5x3-6 


17-0 


— 


1813 


S 


112 


148 


20 


12 


27-6 


19-7 


6-3 


I2-I X4-o 


— 


3'5 


1811 


? 


120 


142 


21 


n 


28-1 


i8-6 


6-i 


11-9x4-6 


16-9 


37 



IN NEW GUINEA, 1932-1949 



277 



Eudromicia caudata (Milne-Edwards) 

Dromicia caudata Milne-Edwards, 1877, C. R. Acad. Sci., Paris, 85: 1 079-1 080. 

Type locality : Arfak Mountains, Dutch New Guinea. 
Eudromicia caudata (Milne-Edwards), Tate & Archbold, 1937, Bull. Amer. Mus. Nat. Hist. 
73: 384-385. 

Fourteen specimens. Eleven from NE. New Guinea: $50.1390, ? 1391, Saiko, 
Bubu River; $1083, $1084, 1085, 1086 (skin only), Tapu, Upper Ramu River 
Plateau ; <$ 1080, 1081, $ 1082, Baiyanka, SE. Bismarck Range ; <£ 1827, Yanka, 
eastern slopes Hagen Range ; $ 1828, Yandara, Bismarck Range ; and three from 
eastern Papua: <J 1077, $ 1078, Enaena, NE. slopes Mt. Simpson; $ 1079, Boneno, 
Mt. Mura. 

The colour of the pelage of all the specimens is very similar and although Nos. 
1390, and 1391 from the Bubu River are larger than the others their measurements 
are very similar to those given by Tate & Archbold (1937) for caudata from Matsika, 
Papua. 

Measurements in mm. (taken in the flesh) : 







* 
















►« 


31 


3| 




i 






1 
t3 









►si 
1 


^3 


3| 

I* 


^2 


"So 




45 S 






<3 




s 


^ 


<o 


b.0 « 


45 « 


</> 


»*» 


"S « 


<o « 




1 


5 


« 
h 


5 


h3 


op 


N -0 


2 2 

>"-< -G 


55 


5 




*•. 


§ 


50.1390 


6* 


97 


163 


18 


18-5 


25-2 


17-6 


5-4 


II-7X3-6 


15-6 


2-4 


4-0 


4'6 


1391 


? 


106 


174 


i8- 5 


20 


25-8 


18-0 


5-2 


II-5X3-9 


15-5 


2-1 


4-0 


4-6 


1083 


6* 


92 


144 


17 


18 


23*5 


15-8 


5-3 


H-5X3-5 


14-6 


2-0 


3*7 


4-2 


1084 


? 


92 


145 


i7'5 


17 


— 


i6-o 


57 


10-5x3-6 


14-4 


2-0 


3'7 


43 


1085 


$ 


99 


138 


17 


19 


23-8 


16-6 


5-6 


12-0x3-7 


14-8 


2-0 


4-0 


4*3 


1080 


6* 


98 


147 


18 


18 


24-2 


i6-o 


5-5 


12-1x3-9 


14-6 





3-9 


4*5 


1081 


6* 


94 


140 


17 


18 


23-2 


15-6 


5-2 


II-2X 


14-4 


2-0 


3'8 


4'3 


1082 


? 


97 


148 


17-5 


17 


23-9 


16-1 


5'3 





14-9 


2-1 


37 


4*4 


1077 


6* 


IOI 


153 


18-5 


20 


22-6 


15-4 


5-o 


II-OX3-4 


13-8 


2-0 


3-6 


4*1 


1078 


9 


92 


143 


17-5 


18 


23-9 


i6-o 


5-4 


II-8X3-7 


14-6 


2-0 


3-8 


4-1 


1079 


? 


95 


148 


18 


19 


24-2 


i6-o 


5-o 


IO-8X3-9 


14-9 


2-0 


4-0 


4'3 


1827 


6* 


108 


145 


19 


18 


25-1 


17-1 


5-i 


II-6X3-7 


15-5 


2-0 


3-9 


4-2 


1828 


$ 


94 


140 


18 


18-5 


24-0 


16-7 


5'4 


II-3X3-7 


— 


2-2 


— 


41 



Dactylopsila trivirgata melampus Thomas 

Dactylopsila melampus Thomas, 1908, Ann. Mag. Nat. Hist. 1: 122. 

Type locality: Kokoda, Mambare River, SE. British New Guinea. 
Dactylopsila hindenburgi Ramme, 1914, S.B. Ges. naturf. Fr. Berl. 1914: 413. 

Type locality: Sattelberg, NE. New Guinea. 
Dactylopsila biedermanni Matschie, 1916, Mitt. Zool. Mus. Berl. 2: 303. 

Type locality : Upper Aroa River, Papua. 
Dactylopsila trivirgata melampus Thomas, Tate & Archbold, 1937, Bull Amer. Mus. Nat. 
Hist. 73: 393- 

In all fifty-five specimens. Thirty-six from NE. New Guinea: twenty of these 
from the Kratke Mts. ; $ 50.1305, juv. $ 1304, ? 1307, 1306, Arau district; <J 1291, 
$ 1292, juv. ?I293, Kambaidam; $ 1302, juv. $ 1301, Apimuri (Buntibasa district) ; 



ZOO. I, 10 



pp 



278 



MAMMALS COLLECTED BY MR. SHAW MAYER 



juv. £ 1303, Yampara; <J 1297, 1296, 1295, 1294, $ 1300, 1299, juv. $ 1298, Bunti- 
basa district; ^1309, 1308, juv. $1310, Kuraka; twelve from near the Upper 
Waria River: $ 1322, Bubu River district; <J 1313, ? 1314, 1318, 1319, 1320, 1321, 
juv. $ 1316, juv. $ 1317, juv. <$ 1311, juv. <J 1312, juv. $ 1315, Garaina; one $ 1014 
from Baiyanka, SE. Bismarck Range ; $ 1822 from Guyebi, Bismarck Range ; and 
three, juv. $ 1819, ad. $ 1820, juv. $ 1821, from Menebe, 8 miles east of Hagen 
Range, Sepik-Wahgi Divide. The following eighteen specimens are all from eastern 
Papua: $1017, juv. $1018, juv. ? 1016, Ikara, NE. slopes Mt. Simpson; £ 1019, 
1020, $ 1022, 1024, 1025, 1026, 1027, 1028, juv. $ 1023, Enaena, NE. slopes Mt. 
Simpson; young ad. <J 1034, J uv - S *035» $ 1036, Boneno, nr. Mt. Mura; (J 1030, 
$ 1031, Wapona, N. slope Maneao Range. 

This excellent collection of skins shows a great deal of variation in the length of 
hair. This is very obvious in the series from the Kratke Mts., which have on the 
whole longer hair and bushier tails than those from other parts of NE. New Guinea 
and eastern Papua, though one or two of these are similarly long haired. It is 
particularly marked in the young specimens. In No. 1301 the hair on the rump 
reaches a maximum for any specimen of about 8 cm. in length. Two specimens, 
juv. (J 1819, ad. $ 1820, collected by Mr. Shaw Mayer in 1946 at Menebe, Sepik- 
Wahgi Divide, NE. New Guinea, 6,000 ft., about 120 miles to the north-east of the 
Kratke Mts., also have long hair and bushy tails. It may be that the length of the 
hair is associated with the age of the animal, as on the whole larger specimens have 
shorter hair. 

The amount of greyish hair in the tail varies from virtually none (Nos. 1310, a 
very dark specimen, 1293, 1294, and 1295, all from the Kratke Mts.) to about two- 
thirds (Nos. 1297 and 1322). Several specimens (mostly females) have white tips to 
their tails (? 1307, 1306, 1299, 1018, young, of black tipped, $ 1017, 1022, 1023, 
1024, 1036, (J 1313, 1020, 1014) 

Specimen No. 1020, an adult (J, is of interest as the black chin spot is divided in 
two by a white stripe running from the throat to the middle of the lower lip, as in 
typical trivirgata or t. infumata. In every other respect it is typical melampus. 

The following are the measurements in millimetres of fourteen adult males and 
twenty-two adult females of melampus : 





Extremes 


A verage 


Standard 
deviation 




a 


? 


<J 


? 


<$ 


? 


Head and body .... 

Tail 

Hind foot ..... 
Ear ...... 


233-287 

263-398 

46-51 

27-31-5 


220-327 

243-382 

44-52 

26-32 


249 
348 

48 
29 


246 

344 

48 

29 


131 

29-3 

1-9 
1-6 


10-3 
25-0 

2-4 
17 



Dactylopsila tatei sp. n. 

Type locality: Mts. above Taibutu village, Faralulu district, West Fergusson 

Island, SE. New Guinea, 2,000-3,000 ft. 
Type: Adult <j> 50.1327, collector's No. 433, 30 July 1935. Skin and skull. 



IN NEW GUINEA, 1932-1949 



279 



Paratypes: $ 50.1325, collector's No. 421, 1326, collector's No. 423, 1329, collector's 
No. 435, juv. 1328, collector's No. 434 (of 1327), 1331, collector's No. 438 (skull 
and piece of skin), $ 1323, collector's No. 425, 1324, collector's No. 431, 1330, 
collector's No. 439 (skull and piece of skin), Mts. above Taibutu village, Faralulu 
district, West Fergusson Island, SE. New Guinea, 2,000-3,000 ft. 

These nine specimens of Dactylopsila taken in West Fergusson Island are most 
nearly allied to D. t. trivirgata. The black and white markings on the head and back 
are very similar, the median stripe being a little darker than the two lateral ones. 
The hairs on the backs of the fore and hind feet are whitish. They are at once 
distinguished from trivirgata by the absence of the large black chin spot and by 
the shorter tail which, on the upper side, has only a few or no grey hairs near the 
base. On the underside the transition from grey to black takes place at about 70-80 
mm. from the base. The tip of the tail is white. 

The skull is very similar to that of trivirgata but is smaller. 

Measurements in mm. of the type and paratypes (taken in the flesh) : 



1 




g 


1 


"0 




^ 


1* 

^ 


3 
►« 


< 


1 


"8 

•2 -~ 


^ 
OQ is 


•"3 
•1 

•1 ? 


■ft, 


„ 


50.1327 Type 
1325 
1326 
1329 
1328 
i33i 
1323 
1324 
1330 




O 
$ 

$* 
O 

5 

c? 


213 

200 

195 
195 
130 

183 
181 
173 

210 


274 

270 
267 
273 
179 

262 

283 

265 

286 


45 
44-5 
43 
43 

33-5 

42 

46 

43-5 

46 


24 
24 

22-5 

24 

20 

23 

22 

22 

24 


4 8-6 
48-4 

47-3 
46-7 
35-o 
44-8 
46-0 
43-6 
48-8 


35-5 
36-0 
35-9 
34-o 
25-0 
3i-8 
33-9 
32-2 
36-4 


7-3 
7-2 
7-5 
7-6 
6-3 
6-8 
7-2 
7-2 
8-o 


17-8x7-8 
I7-9X 8-2 
17-8x7-5 
16-1x6-8 
13-7x6-3 
i6-ox 6-o 
i7-ox 6-7 
15-8x5-7 
17-7x7-2 


25-0 
24-9 
25-2 
25-0 

23-9 
24-5 
23-1 
26-0 


3-0 

2-4 

2-2 

2-5 

2-2 

2-9 
2-4 
2-7 
3-3 


6-2 

6-5 
7-o 

6-2 

6-5 
6-6 
6-6 
7-o 


7-o 
6-7 
7-2 
6-6 

6-7 
7-0 
6-6 
7-3 


n-7 
n-6 
n-8 
n-4 

n-7 

12-0 

n-4 
n-6 


8-2 

8-i 
8-4 
8-i 

8-4 
8-5 
8-i 
8-o 



* juvenile 

Dactylonax palpator (Milne-Edwards) 

Dactylopsila palpator Milne-Edwards, 1888, Mem. Soc. Philom. Centenaire, Paris: 173-177. 

Type locality: Aroa River, Papua. 
Dactylopsila palpator emstmayri Stein, 1932, Z. Sdugetierk. 7: 254. 

Type locality: Junzaing, Saruwaged Range, Huon Peninsula, New Guinea. 
Dactylonax palpator (Milne-Edwards), Tate, 1945, Amer. Mus. Novit., No. 1305:5. 

Twenty-two specimens. Four from eastern Papua: juv. ^50.1032, $1037, ] uv - 
$ 1033, Boneno, Mt. Mura; young ad. ? 102, Enaena, NE. slopes Mt. Simpson; and 
eighteen from NE. New Guinea: £ 1015, Baiyanka, SE. Bismarck Range; $ 1332, 
Kuraka, Kratke Mts.; ^1336-1340, $1341-1345, Bubu River district; $1333- 
1335 (one young in pouch, collectors No. 4860), Saiko, Bubu River; <J 1823, juv. £ 
1824, Yanka, eastern slopes Hagen Range ; $ i825,Tomba, SW. slopes Hagen Range. 

In this excellent series there are five specimens which have a well-developed white 
ring of hair round their wrists (Nos. 1337, 1342, 1037, I 336, and 1033) which is the 
only distinctive character given by Stein (1932) for the race emstmayri which Tate 
(1945) suggests is synonymous with palpator ; this certainly appears to be the case 
in this series. It is noticeable that the males grow to a larger size than the females. 



2 8o MAMMALS COLLECTED BY MR. SHAW MAYER 

Measurements in mm. (taken in the flesh) : 







ts 








1° 


.^ 










1 


* 

% 


S3 







hi 




1 IP 
O .3 


1 « 
^s 

N -0 


-0 


►2 


J 




501015 


6* 


255 


235 


49 


28 


57 


46-2 


9-4 


22-5x8-4 


13-6 


9-3 


1340 


6* 


240 


236 


47 


30 


— 


— 


— 


— 


— 


— 


1332 


6* 


239 


227 


50 


27 


57*9 


46-6 


9-2 


22-5x7-7 


129 


8-6 


1338 


S 


238 


212 


48 


29 


— 


— 


— 


— 


— 


— 


1337 


6* 


232 


213 


45'5 


28 


54-o 


43-7 


9-2 


19-9x6-8 


12-5 


8-6 


1336 


6* 


232 


230 


47 


29 


— 


— 


— 


— 


— 


— 


1032 


6* 


220 


212 


47 


29 


54-8 


4i-3 


8-6 


20-2 X5'7 


135 


9-2 


1037 


? 


235 


205 


50 


30 


55-8 


44'4 


8-9 


21-0x6-9 


13-4 


9-0 


1343 


? 


224 


201 


45-5 


27'5 


— 


— 


— 


— 


— 


— 


1345 


? 


222 


215 


46 


30 


— 


— 


— 


— 


— 


— 


1334 


? 


211 


200 


45 


28 


537 


41-6 


8.9 


20-1 x6-4 


12-7 


8-8 


1344 


? 


208 


194 


43-5 


27 


— 


— 


— 


— 


— 


— 


1342 


$ 


208 


207 


43 


25 


517 


43-9 


8-5 


18-8x6-9 


12-5 


8-5 


1335 


? 


208 


203 


45 


26 


50-0 


43'5 


9-5 


19-6 X 6-2 


12-3 


8-3 


1341 


? 


204 


216 


48 


27-5 


— 


— 


— 





— 


— 


1333 


$ 


207 


198 


42-5 


27 


50-6 


42-4 


9-2 


21-5X6-8 


12-0 


8-2 


1823 


s 


235 


212 


46 


26-5 


52-0 


44-8 


9-4 


2I-OX8-2 


13-0 


8-9 


1825 


? 


I96 


200 


44 


25'5 


45*5 


38-3 


8-4 


I7-9X6-8 


I3-0 


9-0 



Petaurus breviceps papuanus Thomas 

Petaurus breviceps var. papuanus Thomas, 1888, Catalogue of the Marsupialia and Monotremata 

in the British Museum: 158. 
Type locality : Huon Gulf, eastern New Guinea. 
Petaurus (Petaurella) papuanus papuanus Tate & Archbold, 1937, Bull. Amer. Mus. Nat. 

Hist. 73: 387. 
Petaurus breviceps papuanus Thomas, Tate, 1945, Amer. Mus. Novit., No. 1305:9. 

Five specimens. Three, <J 50.1377, 1378, $ 1379, from Taibutu, Faraluhi district 
West Fergusson Island, SE. New Guinea; and two from NE. New Guinea: <£ x 38o, 
Garaina, Upper Waria River; $1826 (white-tipped tail), Degabaga, 8 miles east 
Hagen Range, Sepik-Waghi Divide. 

Petaurus breviceps tafa Tate & Archbold 

Petaurus (Petaurella) papuanus tafa Tate & Archbold, 1935, Amer. Mus. Novit., No. 810: 1; 
1937, Bull. Amer. Mus. Nat. Hist. 73: 387. 
Type locality: Mt. Tafa, Central Division of Papua. 
Petaurus breviceps tafa Thomas, Tate, 1945, Amer. Mus. Novit., No. 1305:10. 

Ten specimens. Seven from NE. New Guinea: (£50.1382, 1381, $1383, Kambaidam, 
Kratke Mts. ; (£1385, 1384, $1386, Saiko, Bubu River; (£1069, Baiyanka, SE. 
Bismarck Range; and three from eastern Papua; 3 ^oyo, Enaena, NE. slopes Mt. 
Simpson; 3 1071, $ 1072, Boneno, Mt. Mura. 

These highland specimens are new to our collection. They were taken between 
4,000 and 7,500 ft. and agree with Tate & Archbold's description of the dark- 
coloured mountain race tafa (especially when comparing teeth measurements), 



IN NEW GUINEA, 1932-1949 281 

though the pelage on the back is only 9-10 mm. long as compared with 12 mm. in 
the type. They are smaller than typical papuanus, grey ventrally with only a slight 
buffy overwash. Three of them, <J 1382, $ 1386 and £ 1072, have a patch of buffy 
hairs in the middle of the belly, and $ 1069 nas a white tip to its tail. 

Measurements of four specimens which range from the smallest to the largest 
are given. 

Measurements in mm. (taken in the flesh) : 





















d 























.0 








■JS 






i 




•IS 

"3> 














la 








"o 
13 




a 




.0 
O * 


.0 


"3 


3* 


■ft, 

5 i 


„ 












CO 


































4 


£ 


s 


53 


4 


£ 




s Si 


03 


1 


« 




■ft, 


5 


ft, 


g 


^ 


50.1072 


¥ 


123 


154 


22 


23 


27-6 


22-4 


67 


16-2 


io-7X 5-0 


16-3 


i-8 


6-6 


4-5 


i-o 


i-gx i-8 


1-5x1-5 


1381 


3 


128 


178 


23 


22-5 


31-0 


23-3 


7-1 


15-7 


13-4x5-8 


17-7 


1-9 


7-2 


4-7 


1-4 


2-ox i-8 


i-6x 1-7 


1382 


3 


133 


160 


22-5 


22 


30-9 


24-2 


7-0 


i6-o 


12-9x5-3 


17-7 


1-9 


73 


4-8 


i-3 


I-9X i-8 


i-6x 1-7 


1385 


<J 


137 


174 


24 


23-5 


30-9 


23-0 


7-2 


16-2 


I3-4X5-8 


17-9 


1-9 


7-1 


4-7 


i-4 


i-8x 1-7 


I-5X i-6 



Pseudocheirus (Pseudochirops) cupreus cupreus Thomas 

Pseudochirus cupreus Thomas, 1897, Ann. Mus. Stov. nat. Genova, 38: 145-146. 

Type locality : Mount Owen Stanley, British New Guinea. 
Pseudochirus (Pseudochirops) cupreus obscurior Tate & Archbold, 1935, Amer. Mus. Novit., 

No. 810: 3-4. 
Pseudocheirus (Pseudochirops) cupreus cupreus Thomas, Tate, 1945, Amer. Mus, Novit., No. 

1287: 20-21. 

Seventeen specimens, all from NE. New Guinea: ^50.1369, 1370, 1372, juv. 
^1368, juv. c? 1371, Saiko, Bubu River; $ 1373, I374> 1375, Bubu River 
district ; <J 1367, Sasara, Kratke Mts. ; (J 1066, juv. £ 1067, ? 1068, Baiyanka, SE. 
Bismarck Range; ^1063, juv. ^1062, $1065, juv. $1064, Tapu, Upper Ramu 
River Plateau ; $ 1818, Yanka, eastern slopes Hagen Range. 

The pelage of these specimens does not differ markedly from that of the adult 
type specimen, but the immature specimens are much darker. 

They extend the range of this species to the north-west as far as the Upper Ramu 
River Plateau. 



Pseudocheirus mayeri Rothschild & Dollman 

Pseudochirus mayeri Rothschild & Dollman, 1932 (November) Abstr., Proc. Zool. Soc. Lond. 
No. 353:i5- 

Type locality: The Gebroeders, Weyland Mts., Dutch New Guinea. 
Pseudochirulus pygmaeus Stein, 1932 (December), Z. Sdugetierk. 7: 257. 

Type locality: Sumuriberg, Weyland Mts., Dutch New Guinea. 

One specimen, <J 50.1808, Tomba, SW. slopes Hagen Range, NE. New Guinea. 

This species is found high up in the mountains between 6,000 and 12,000 ft. It 
appears that this is the first record of its occurrence outside Dutch New Guinea and 
so extends its known range into eastern New Guinea. The pelage is dense, and very 
soft, the upper parts greyish brown and the underparts buff with the bases of the 
hairs grey. The hands and feet are a light brownish buff and there is a whitish patch 






282 



MAMMALS COLLECTED BY MR. SHAW MAYER 



of hairs at the base of the ears. The measurements of this specimen agree very 
closely with those for mayeri (pygmaeus) given by Tate (1945). 
Measurements in mm. (taken in the flesh) : 







?s 





































<s 




















* 

^ 




"o 




-0 


,w 


















































«, 
ts 




^s 


g§ 


03 




"g 








CO 


*s 






v 


s &* 


&o ^ 














fe; 


^ 

m 


« 
^ 


&} 


« 

^ 


O s 
0^ 




£ 


3*1 


■fc, 


§ 


§ 


"s 


50-1808 


<J 


194 


176 


25 


20 


42*2 


24-2 


13-5x6-0 


2-ix 1-5 


n-7 


7-5 


2-7X2-1 


2-2X 1-8 



Pseudocheirus forbesi forbesi Thomas 

Pseudochirus forbesi Thomas, 1887, Ann. Mag. Nat. Hist. 19: 146. 

Type locality: Sogere, Astrolabe Range, SE. New Guinea, 2,000 ft. 
Pseudocheirus forbesi forbesi Thomas, Tate, 1945, Amer. Mus. Novit., No. 1287:11. 

Five specimens all from eastern Papua: ^50.1039, 1040, juv. ^1042, 2 1041, 
Enaena, NE. slope Mt. Simpson; § 1038, Boneno, Mt. Mura. 
Measurements in mm. (taken in the flesh) : 
















"e 























-0 


/o 



































<s 




^ 




•~~. 


1 -« 








3 




^s 




"« 




£* 


§ ^ 


S3 


~t~ 










s 


>< 
e 
hi 


1 £ 


«10 <s 

N -0 




a. 


it 


50.1039 


s 


250 


238 


36 


20 


49-8 


29-2 


17-8X7-0 


14-2 


8-9 


1040 


6* 


277 


271 


37 


19 


52-7 


29-9 


— X7-8 


141 


9-2 


1041 


¥ 


227 


238 


33 


19 


457 


27-2 


16-1 x6-7 


13-5 


8-8 


1038 


$ 


230 


255 


33 


20 


47'4 


28-0 


— X6-6 


13-8 


8-8 



Pseudocheirus forbesi larvatus (Forster & Rothschild) 

Phalanger larvatus Forster & Rothschild, 1911, Ann. Mag. Nat. Hist. 7: 1337. 

Type locality: Rawlinson Mountains, Huon Peninsula, New Guinea. 
Pseudochirulus capistratus Matschei, 1915, S.B. Ges. naturf. Fr. Berl. 1915: 92. 

Type locality: Schrader Mts. between the Sepik and Ramu Rivers, NE. New Guinea. 
Pseudochirulus barbatus Matschie, 1915, ibid. : 93. 

Type locality: Sattelburg, north of Huon Gulf, NE. New Guinea. 
Pseudocheirus forbesi larvatus (Forster & Rothschild), Tate, 1945, Amer. Mus. Novit., No. 
1287:12. 

Fourteen specimens all from NE. New Guinea: <$ 50.1346, juv. 6* I 347> ? sex 1348 
and 1349 (skulls only), Kambaidam, Kratke Mts. ; <J 1350, Buntibasa, Kratke Mts. ; 
$ 1351, Kuraka, Kratke Mts. ; $ 1043, $ 1044, Tapu, Upper Ramu River Plateau; 
S 1045, Baiyanka, SE. Bismarck Range; ^1354, J 35 2 > I 353> I 355> Saiko, Bubu 
River ; $ 1809, Yanka, eastern slopes Hagen Range. 

There is a great similarity between the general colouring of these specimens and 



IN NEW GUINEA, 1932-1949 



283 



P. f. forbesi, but they are larger (see measurements) and usually have much darker 
tails. 

Measurements in mm. (taken in the flesh) : 



5< 


CO 


Head and body 


'1 


"0 




Condylo-basal 
length 


Zygomatic 
breadth 


5S 


J 


M 


50.1346 


0* 


295 


300 


39 


20 


58-9 


357 


2I-OX8-4 


15-3 


IO-2 


1343 


? 


— 


— 


— 


— 


54-9 


3i-3 


— 7'3 


15-9 


10-4 


1349 


? 


— 


— 


— 


— 


53-2 


29-7 


— 6-8 


15-0 


9-9 


1350 


6* 


— 


— 


39 


20 


56-0 


30-6 


20-4x6-8 


i5'9 


10-5 


I35i 


9 


270 


285 


38 


19 


57-2 


3i-i 


20-9x7-5 


i5'5 


io-o 


1043 


6* 


292 


280 


35 


20 


58-8 


34-7 


20-8 x 7-0 


i5-i 


9-8 


1044 


¥ 


236 


250 


31 


19 


50-6 


28-5 


— 6-5 


14-7 


9-9 


I045 


3 


298 


310 


40 


20 


59-o 


32-8 


~ 7'5 


16-2 


10-9 


1354 


(J 


286 


302 


42 


23-5 


56-4 


32-1 


21-4x8-1 


15-4 


IO-I 


1352 


3 


260 


270 


38 


21-5 


52-1 


29-2 


l8-2 X 7-0 


14-8 


9-9 


1353 


«J 


298 


319 


41 


23-5 


56-7 


32-7 


19-8x8-8 


15-2 


9-9 


1355 


6* 


323 


307 


39 


23 


61-7 


34-6 


22-0x9-8 


15-0 


9-5 


1809 


0* 


251 


268 


40 


19 


52-7 


307 


18-9x7-6 


15-6 


io-6 



Pseudocheirus canescens gyrator Thomas 

Pseudochirus canescens gyrator Thomas, 1904, Ann. Mag. Nat. Hist. 14: 401. 

Type locality, Lindum Creek, Gira River district, NE. of the Central Range, Dutch New 
Guinea, 600 ft. 
Pseudocheirus canescens gyrator Thomas, Tate, 1945, Amer. Mus. Novit., No. 1287: 14-15. 

Four specimens all from eastern Papua: ^50.1046, $1048, 1047, Enaena, NE. 
slopes Mt. Simpson ; $ 1049, Ikara, NE. slopes Mt. Simpson. 

These four specimens from eastern Papua are very similar to the type specimen, 
which up to now appears to have been unique. The general colour is brownish-grey 
(two specimens, Nos. 1046 and 1047, are rather greyer than the others) ; the head, 
face, cheeks, fore and hind limbs are pale brown ; and there is a well-marked fuscous 
frontal stripe and a darker line down the middle of the back. The ears are fuscous 
and have dark hairs round their bases. Ventrally the hairs are brownish-buff, their 
bases sometimes grey. 

Measurements in mm. (taken in the flesh) : 



1 

fe; 










ts 




s 




'S 

too « 

^ Si 

N -0 


1! 


00 


"So 






n 


50.1047 
1046 
1049 
1048 


? 

9 

? 


201 
225 

230 
234 


173 

192 
200 

182 


29 

33 

28 
28 


17 
18 
16-5 
17 


447 
46-9 

47-3 
47'3 


26-7 
27'5 
27-5 
28-0 


21-8 
22-6 
22-1 
23'4 


15-8x5-7 
15-9x6-7 
17-8x6-0 
16-9x6-8 


22-8 
23-7 
24-5 
24-4 


3-9 
4-3 
4'3 

4-7 


12-5 
12-3 

12-2 

u-7 


8-o 
7 -8 
7-8 
7-7 



28 4 



MAMMALS COLLECTED BY MR. SHAW MAYER 



Pseudocheirus (Pseudochirops) corinnae corinnae Thomas 

Pseudochirus corinnae Thomas, 1897, Ann. Mus. Stor. nat. Genova, 38: 142-144. 

Type locality : Mountains of Vanapa River district, British New Guinea. 
Pseudocheirus {Pseudochirops) corinnae corinnae Thomas, Tate, 1945, Amer. Mus. Novit., No. 

1287: 20. 

Fourteen specimens all from NE. New Guinea: (J 50.1050, $ 105 1, Baiyanka, SE. 
Bismarck Range ; $ 1052, Tapu, Upper Ramu River Plateau ; <J 1358, 1359, 1360, 
$ 1361, Buntibasa district, Kratke Mts. ; $ 1356, 1357, Kuraka, Kratke Mts. ; $ 1362, 
$ 1363, 1364, 1365, Saiko, Bubu River district ; $ 1366, Bubu River district. 

These specimens closely resemble the co-type and other specimens of corinnae in 
this Museum except for No. 136 1, an adult female in which the dorsal pelage re- 
sembles that of P. c. argenteus. It is a bright rusty colour especially on the rump 
and tail, but ventrally is just the same colour as corinnae, a light dirty yellowish- 
grey. 

These specimens extend the range of the species to the north-west as far as the 
Upper Ramu River Plateau. 

Measurements in mm. (taken in the flesh) : 







S 














-« 


H 

n 



















►ss 






1 








*< 


h 


t3 

53 


r"** 


"0 


x 


.3 


^ 


"q 


.3 

3 




\ 




s 


<» 


§ 




ts 


« 


5s $> 


<s 


«a 


s ^ 


1 




fe; 


$ 


tq 


h 


5 


^ 


oq 


^£ 


£ 


»H 


^ ^ 


■^ 


§ 


50.1052 


6* 


340 


294 


42 


24 


617 


397 


21-0x9-9 


36-3 


4-9 


19-8 


13-0 


1050 


6* 


318 


288 


49 


22 


57-2 


36-i 


21-2X8-6 


33-8 


4'4 


20-0 


I3'6 


1051 


? 


300 


315 


47 


23 


56-6 


37'2 


20-3 X9-o 


327 


5-6 


20'7 


12-9 


1356 


$ 


325 


315 


44 


23 


6o-i 


37-8 


21-5 xg-o 


35-3 


5-o 


20-8 


13-8 


1358 


6* 


340 


345 


50 


25 


63-2 


42-3 


22-OX II-O 


367 


5-i 


20-5 


13-3 


1357 


$ 


317 


305 


47 


22-5 


59-4 


37'5 





35-2 


5-2 


20-0 


13-2 


1363 


? 


340 


335 


5i 


24 


62-3 


40-0 


20-0X9-8 


36-4 


6-2 


19-5 


12-8 


1364 


? 


350 


320 


47-5 


25 


62-4 


40-0 


22-2 X IO-6 


36-6 


5-6 


19-6 


12-7 


1365 


? 


315 


315 


46-5 


25 


— 


38-4 


c. 20-0x9-2 


— 


5-4 


20-2 


13-4 


1362 


6* 


325 


305 


48 


24 


62-1 


40'3 


21-7 X97 


36-9 


5-o 


20-7 


137 


1366 


? 


345 


323 


47-5 


26 


61-4 


39-o 


2I-OX IO-8 


36-4 


5'4 


19-8 


12-8 



Pseudocheirus [Pseudochirops) corinnae fuscus subsp. n. 

Type locality: Ikara, NE. slopes Mt. Simpson, eastern Papua, SE. New Guinea, 
±4,000 ft. 

Type: Adult $ 50.1058, collector's No. 761, 16 August 1940. Skin and skull. 

Paratypes: $ 50.1054, collector's No. 763, 1055, collector's No. 764, young ad. 1053, 
collector's No. 762, $ 1057, collector's No. 757, young ad. 1056, collector's No. 753, 
Ikara, NE. slopes Mt. Simpson, eastern Papua, ±4,000 ft. ; $ 1059, collector's 
No. 774, 1060, collector's No. 785, Enaena, NE. slopes Mt. Simpson, eastern 
Papua, ± 5,000 ft. ; $ 1061, collector's No. 945, Boneno, nr. Mt. Mura, eastern 
Papua, 4,000-5,000 ft. 

Rather darker than typical corinnae. Most of the hairs are a dark greyish-brown 



IN NEW GUINEA, 1932-1949 285 

tipped with silver, buffy, rust, or black. The black median dorsal stripe is well 
defined and there is an indistinct dark brown stripe along each side of the back. 
There is a patch of white or yellowish-white hairs at the base of the ears. 

This race can be at once distinguished from typical corinnae by the very distinct, 
usually diamond-shaped patch of white hairs on the throat and chest. The rest of 
the ventral surface is a rather dark yellowish- grey or yellowish-brown. 

The general appearance and the measurements of the skull are very similar to 
those of typical corinnae; one of the main differences, however, is in the length of 
p 4 -m 4 and m 1-3 . Both are usually longer, the greatest lengths recorded for speci- 
mens in this collection being p 4 -m 4 = 21-8 with m 1-3 = 14-3, compared with maxi- 
mum lengths for typical corinnae of p 4 -m 4 = 20-8 with m 1-3 = 13-8 (see table). 
The frontals are often more depressed than those of typical corinnae, the supra- 
orbital ridges are well developed, and in adult specimens there is a well-developed 
sagittal crest. 

Measurements in mm. of the type and paratypes (taken in the flesh) : 







£ 














-si 


41 












-0 








•^5 






"§> 










00 









^ 


1 

.3 

00 


^ 




Si 


i 1 


§ 


n 


1 






S 




e 

cq 




1 


1 


s *< 

^ 


1 


§ 


5O.IO56 


? 


333 


329 


47 


25 


62*0 


38-0 


22-0 X 9"8 


36-4 


5-9 


21-5 


13-8 


I057 


? 


330 


35o 


5i 


25 


62-8 


39-8 


20-5X9-8 


37.0 


5'4 


21-3 


14-4 


1058 Type 


? 


352 


310 


49 


25 


6i«4 


39-i 


2I-8XIO-I 


35-5 


5-8 


20-5 


13-4 


1053 


tJ 


334 


326 


— 


27 


6i-o 


40-0 


19-7 X io-6 


36-7 


5'2 


21-4 


I3'9 


1054 


6* 


337 


345 


50 


27 


64-0 


42-8 


22-9 X IO-8 


37-6 


5'4 


21-8 


14*3 


1055 


a* 


354 


37i 


5i 


27 


65-2 


44-7 


22-1 X 11*5 


38-3 


5-7 


21-4 


13-7 


1059 


? 


326 


335 


50 


25 


58-8 


37-6 


— X 87 


35-o 


5-4 


21-5 


14-6 


1060 


? 


319 


315 


49 


28 


58-7 


38-2 


2I-OX IO-5 


34-8 


4-8 


21-5 


13-8 


1061 


$ 


339 


354 


50 


26 


62-4 


40-5 


21-2 X IO-O 


36-9 


5'4 


20-9 


I3'7 



Phalanger orientalis orientalis (Pallas) 

Didelphis orientalis Pallas, 1766, Miscellanea Zoological 59-62. 

Type locality : Amboina, off SW. coast of Ceram. 
? Phalangista quoy Gaimard, 1824, Bull. Sci. Nat. Paris, 1: 271. 

Quoy & Gaimard, Voyage. . . . Uranie et Physicienne. Zoologie: 58, t.l. Waigeu. 
ICoescoes amboinensis Lac6pede, 1801, Mem. Inst. Paris, 3: 491, t.l. Amboina. 
Phalanger 0. orientalis (Pallas), Tate, 1945, Amer. Mus. Novit., No. 1283: 1-31. 

Three specimens, ^50.1285, 1286, juv. 1287 (skull and piece of skin) from the 
Faralulu district, West Fergusson Island, SE. New Guinea. 

These specimens are very similar to others in the British Museum's collection. 
The general colour is white, tinged, especially on the throat and sides of the neck, 
with yellow. The dorsal surface is covered with longer black-brown hairs which 
are most numerous on the head and neck, on all four feet and base of tail, and, of 
course, along the dorsal line, which is well defined. 

zoo. 1, 10 q q 



286 MAMMALS COLLECTED BY MR. SHAW MAYER 

Measurements in mm. (taken in the flesh) : 







§ 

































O 




"5 

"So 

<4i 


_y> 


1 












<» 




« 




^ 






ig 


'£ ^ 






























■* 








s 








^ 
1 

3 






1 ^ 

N .5 


^ -O 


<3 


J 

"Oh 


"g 


^ 


s 


50.1285 


a 


389 


362 


60 


25-5 


75-5 


50-9 


40-0 


29-3 X 12-8 


22-2 


13-6 


47 


4-6x3-8 


1286 


a 


397 


340 


— 


— 


76-1 


57-5 


4 2-7 


34-8x14-6 


22-8 


15-0 


4'4 


5-0x4-0 



Phalanger vestitus (Milne-Edwards) 

Cuscus vestitus Milne-Edwards, 1877, C.R. Acad. Sci. Paris, 85: 1080. 

Type locality: Karons Mountains, Tamrau Mountains, northern Vogelkop. 
Phalanger carmelitae Thomas, 1898, Ann. Mus. Stor. nat. Genova, 39: 5. 

Type locality : Upper Vanapa River, British New Guinea. 
Phalanger sericeus Thomas, 1907, Ann. Mag. Nat. Hist. 20: 74. 

Type locality: Owgarra, Angabunga River, SE. New Guinea. 
Phalanger coccygis Thomas, 1922, Ann. Mag. Nai. Hist. 9: 673. 

Type locality: Surawaged Mts., Huon Peninsula, New Guinea. 
Phalanger vestitus (Milne-Edwards), Tate, 1945, Amer. Mus. Novit., No. 1283: 16. 

Forty-six specimens. Thirty-two from NE. New Guinea: $ 50.1269, 1270, $ 1271, 
1272, 1273, juv. $ 1274, Sasara, Kratke Mts. ; $ 1275, juv. ? 1276, Buntibasa district, 
Kratke Mts.; $ 1277, I2 7^, 1279, $ 1280, juv. $ 1281, Saiko, Bubu River; $ 1262, 
1263, 1264, 1265, $ 1266, 1267, 1268, 1284, Bubu River district ; juv. $ 1283, Arau, 
Kratke Mts. ; $ 992, 996, duplicate — collector's No. 735 (skull only), § 997, juv. $ 1000, 
juv. $ 998, juv. S 999> Baiyanka, Bismarck Range; $ 993, $ 994, juv. $ 995, Tapu, 
Upper Ramu River Plateau; <$i8ij, Yanka, eastern slopes Hagen Range; and 
fourteen from eastern Papua; ^989, 990, Mt. Mura; ^991, 1002, $1004, juv. 
$ 1003, Boneno, Mt. Mura; $ 1001, Bibitau, Mt. Orian; $ 1005, juv. $$ 1009 (young 
of 1005), 1008, 1006, 1010, juv. (J 1007, Enaena, NE. slopes Mt. Simpson. 

This excellent collection of skins which includes those of fourteen juvenile speci- 
mens indicates the great variability in the colour and length of hair in this species, 
from a short-haired pale silvery-brown specimen (adult $ 1004) through intermediate 
forms which are nearer the type specimen of carmelitae Thomas (1898) to a dark 
brown longer-haired specimen (adult $ 994) which is very similar to the type of 
sericeus Thomas (1907). The type of coccygis Thomas (1922) also fits into this 
series, supporting the view suggested by Tate (1945) that carmelitae, sericeus, and 
coccygis are synonymous. There is also a great similarity in appearance between the 
juvenile forms and a specimen of P. vestitus which we have in this Museum and which 
I am unable to distinguish from the young specimens in the Shaw Mayer collection. 
The pelage of these specimens is usually longer than that of the adults, and is a 
little darker and somewhat grizzled, especially along the sides of the body ; and the 
dark brown mid-dorsal line is more clearly defined. 

The type of vestitus [Cuscus vestitus Milne-Edwards), which I have not seen, is 
a young specimen and the description of it agrees with that of the young specimens 



IN NEW GUINEA, 1932-1949 



287 



in this collection. It appears, therefore, that carmelitae, sericeus, and coccygis are 
synonymous with vestitus and not races of it as suggested by Tate (1945). 

The hair on the backs of specimens 1273 and 1284 is very short as it has been 
clawed off by the young. 

The following are the measurements in millimetres of seventeen adult males and 
fourteen adult females of vestitus : 











Standard 




Extremes 


A verage 


deviation 




3 


$ 


6* 


$ 


6* 


? 


Head and body 


327-437 


353-455 


408 


407 


28-1 


29-3 


Tail 








305-404 


333-387 


356 


367 


26-3 


15-2 


Hind foot 








56-66 


57-62 


60 


60 


3-9 


2-0 


Ear 








19-27 


20-26 


23 


24 


2-1 


2-1 


Basal length 








68-3-78-8 


68-8-78-4 


74*4 


73-i 


3-3 


2-9 


Zygomatic breadth 








47-0-56-9 


45-4-52-3 


5i-i 


48-9 


2-6 


1-8 


Mastoidal breadth 








37'4-45-3 


37-0-46-4 


41-8 


41-0 


2-4 


2-5 


Nasals, length . 








26-3-31-9 


25-6-31-3 


29-0 


27-7 


1-9 


1-9 


Nasals, breadth 








10-0-14-5 


10-1-13-1 


12-3 


u-9 


i-i 


0-9 


p 4 -m 4 








23-7-26-8 


23-7-26-7 


25-2 


25-0 


0-9 


0-9 


m 1 -* 








I57-I77 


15-2-17-9 


16-6 


16-3 


o-5 


0-7 



Measurements in mm. of twelve juvenile specimens (taken in the flesh) : 



5s 

1 


* 

<» 
^ 




■-0 


Hind foot 


3 




do « 

N .0 




•^2 

to 

1 


50.1274 


? 


275 


270 


46 


20 


53-5 


36-8 


29-7 


23-1 x io-8 


1276 


2 


325 


295 


53 


21 


60-7 


40-0 


32-6 


24-2 X IO-I 


I28l 


¥ 


315 


300 


49 


23 


59'i 


39-i 


31-0 


20-OX II-5 


1283 


? 


280 


300 


51 


20 


54-9 


38-0 


3i-5 


22-9 X IO-O 


IOO8 


? 


328 


342 


54 


24 


6o-6 


38-5 


3i-i 


2I-7XII-I 


IOO6 


$ 


355 


323 


— 


— 


63-4 


42-0 


34-i 


25-0x10-5 


IOIO 


¥ 


34i 


341 


57 


26 


64-7 


40'5 


33-2 


25-0x10-3 


1007 


s 


406 


392 


63 


27 


73-4 


48-0 


38-8 


26-0 X 12-0 


998 


6* 


343 


340 


57 


23 


64-3 


42-2 


36-2 


XII'7 


999 


s 


364 


323 


59 


24-5 


67-0 


43-2 


36-8 


24-5X11-4 


995 


$ 


338 


335 


52 


21 


60-7 


40-5 


33-6 


22-OX IO-9 


1003 


V 


376 


344 


57 


25 


66-9 


42-0 


33-7 


25-0 X 12-2 



Phalanger gymnotis (Peters & Doria) 

Phalangista gymnotis Peters & Doria, 1875, Ann. Mus. Stor. nat. Genova, 37: 543 



Type locality : Aru Islands. 
Phalanger leucippus Thomas, 1898, ibid. 39: 7-8. 

Type locality : Upper Vanapa River, British New Guinea. 
Phalanger gymnotis (Peters & Doria), Tate, 1942, Amer. Mus. Novit. 



No. 1283: 1-31. 



Ten specimens. Seven from NE. New Guinea: juv. $ 50.1261, Saiko, Bubu River; 
$ 1258, juv. $ 1259, J uv - ? 1260, Bubu River district ; ad. $ 1257, Kambaidam, 



2 88 



MAMMALS COLLECTED BY MR. SHAW MAYER 



Kratke Mts. ; ad. $984, 985, Baiyanka, SE. Bismarck Range; and three from 
Eastern Papua: juv. $986, Enaena, NE. slopes Mt. Simpson; $988 (Boneno Camp), 
Mt. Maneao ; <$ 987, Mt. Mura. 

These specimens extend the range of this species to the north and south of its 
previously recorded range, from the Bismarck Range, NE. New Guinea, to Mt. 
Simpson in the south of eastern Papua. 

The skins show little variation in colour from the type of leucippus (— gymnotis) 
in the British Museum's collection ; the younger specimens are darker, and have a 
more pronounced median dorsal stripe, than the adult specimens. 

Measurements in mm. (taken in the flesh) : 







4S- 


























-0 








*g 
















ts 








* 


/o 


« 








%» 












<a 




^s 








<a 




e 




^ 






is 


'£ ■** 


























■* 




s 

1 




t3 




■^ 




53 


^ 


"3 *3 


s 


§2 




CO 


8 

£3 






^ 

e 

^ 










J 
•ft, 


"s 


50.984 


$ 


415 


318 


64 


27 


7°'3 


48-9 


41-2 


39-8x11-9 


— 


— 


985 


¥ 


4OO 


3IO 


— 


29 


67-8 


45-o 


39-o 


28-8 x 10-9 


— 


— 


988 


s 


472 


355 


65 


30 


80-5 


58-5 


49-1 


34-4X12-9 


25-6 


15-7 


987 


6* 


470 


330 


69 


30 


79-9 


6o-i 


47-6 


33-7x12-6 


25-5 


15-5 


1258 


s 


44O 


320 


63 


28 


78-4 


54'4 


48-1 


34-1X13-6 


24-0 


14-6 


1257 


6* 


465 


35o 


66 


30 


83-1 


6o-i 


50-0 


34-2x15-0 


2 4 -S 


15-0 



Echymipera oriomo Tate & Archbold 

Echymipera oriomo Tate & Archbold, 1936, Amer. Mus. Novit., No. 823:1. 
Type locality: Dogwa, Oriomo River, Western Division of Papua. 

One specimen $ 50.1139, from Tapu, Upper Ramu River Plateau, NE. New 
Guinea. 

One fully adult (old) specimen, judging from the skull of a species which is new 
to our collection. The teeth are very worn down both in the upper and lower jaws, 
and many of them are missing. The tail has been broken off at the root. Miklouho- 
Maclay (1884) mentions that specimens of bandicoots sometimes have the tail lost 
(or bitten off?). 

It is a small-sized species with spinous pelage and relatively small-sized teeth. The 
colour of the pelage agrees with that described for oriomo Tate (1936), and though 
many of the measurements of this specimen are larger than those of the type this 
may be due to differences in age, which may also account for the difference in the 
size of the posterior palatal openings and those of the type oriomo. In the type these 
extend from the front of m 1 to the back of m 2 , whereas in this specimen they extend 
from the front of m 1 to between m 3 and m 4 . Differences in the breadth of the teeth 
(they are wider in our specimen) may be due to wear. 

Measurements in mm. (taken in the flesh ; measurements of type in parentheses) : 
Head and body 291 (244) ; tail — broken off; hind foot 48 (47) ; ear 23-5 ; skull, basal 
length 57-3 (52-3); zygomatic breadth 25-9 (24-0) ; nasals 26-9x5*0 (damaged) 
(24-3x4-8); palatal length 38-6 (35-2); anterior palatal foramina 6-o; posterior 
palatal foramina 8-4 (from front of m 1 to between m 3 and m 4 ) ; teeth (crowns) m 1-3 



IN NEW GUINEA, 1932-1949 289 

9-9 (10-5) ; m 1-2 c. 6-2 ; m 1 (length x breadth) 3-0 x 3-0 ; m 2 missing ; m 3 3-5 X 4- o ; m 1 " 2 
(to front of m 3 ) 6-5. 

Echymipera dor ey ana dor ey ana (Quoy & Gaimard) 

Perameles doreyana Quoy & Gaimard, 1830, Voyage de la corvette V Astrolabe, Zool. 1 :ioo. 

Type locality: Dorey (nr. Manokwari) , Dutch New Guinea. 
Perameles cockerelli Ramsay, 1877, Proc. Linn. Soc. N.S.W. 1: 310, 378. 

Type locality: New Ireland. 
Perameles myoides Gunther, 1883, Ann. Mag. Nat. Hist. 11: 247. 

Type locality : New Britain. 
Br achy metis gar agassi Miklouho-Maclay, 1884, Proc. Linn. Soc. N.S.W. 9: 713. 

Type locality: Maclay Coast (Cape Croisilles to Cape King William), NE. New Guinea. 
Anuromeles rufiventris Heller, 1897, Abh. zool. anthrop. ethn Mus. Dresden, 6:5. 

Type locality: Bongu, Astrolabe Gulf, New Guinea. 
Suillomeles hispida Allen & Barbour, 1909, Proc. New England zool. CI. 4: 44. 

Type locality: Manokwari, Dore Bay, Dutch New Guinea. 
Perameles doreyana and breviceps Cohn, 1910, Zool. Anz. 35: 724. 
Echymipera doreyana doreyana (Quoy & Gaimard) Tate, 1948, Bull. Amer. Mus. Nat. Hist. 

92: 332-333. 

Two specimens: $50.1420, Taibutu, Faralulu district, West Fergusson Island, 
SE. New Guinea ; and $ 138, Wapona, north slope Maneao Range, eastern Papua. 
Measurements in mm. (taken in the flesh) : 







$ 
















T§ 




























-e 










a 


















1 









60 
s 


^ 




S 


*, 

^ 8 


■ft, 














1 




«*, 


1 


V. 


1 


§3 « 


"« 


2 


s 


•2 .§ 

is s 


T 


B 


<M 










































fe; 


W) 


% 


hi 


&3 


&3 


cq 


N .0 


fei 


ft, 


^ £, 


^ 


•ft, 


s 


§ 


S 


a 


£ 


50.1420 


9 


321 


99 


55-5 


27 


62-6 


25-9 


34-6x4-6 


41-0 


4-9 


6-o 


15-6 


n-o 


7-3 


3-9x2-6 


3-5X2-9 


3-9x3-4 


1138 


<J 


342 


84 


65 


32 


69-0 


27-3 


34-7X 6-6 


45-7 


8-7 


6-7 


i8-i 


12-4 


8-i 


4-4 X 2-6 


4-0x3-2 


4-4x4-0 



Peroryctes raffrayanus r affray anus (Milne-Edwards) 

Perameles raffrayanus Milne-Edwards, 1878, Ann. Sci. Nat. Zool. 7: Art. 11: 1-2. 

Type locality: Amberbaki, Vogelkop, Dutch New Guinea. 
Peroryctes raffrayanus raffrayanus (Milne-Edwards), Tate, 1948, Bull. Amer. Mus. Nat. Hist. 
92: 327. 



Five specimens. Four specimens from NE. 
Kratke Mts. ; ? 1411, Sasara, Kratke Mts. ; juv. 
skin), Kambaidam, Kratke Mts.; and one juv. 
Mt. Simpson, eastern Papua. 

Measurements in mm. (taken in the flesh) : 



New Guinea, <$ 50.1407, Kuraka, 
$ 1408, $ 1410 (skull and piece of 
$ 1137, from Enaena, NE. slopes 



9 
"2 




t 
1 


1 


1 

a 


<3 


a 


is 


1 


5 
a 

1 


■2 -S 
11 


3s 
■ex, 
|| 

II 


a 


« 


c. 


1 


°a 




50.1407 
1411 


? 


358 
333 


197 
174 


80 

72 


32 

31-5 


77-4 
72-1 


31-1 
30- 1 


38-6x6-3 
34-9x5-0 


49-7 
47-o 


9-8 

7-5 


n-i 
9-7 


17-2 
16-2 


12-0 

n-5 


8-2 
7-9 


4-3X2-7 
4-0x2-7 


3-8x3-1 
3-8x3-0 


3-9X3-4 

3-7X3-4 



:go 



MAMMALS COLLECTED BY MR. SHAW MAYER 



Peroryctes longicauda ornata (Thomas) 

Perameles ornata Thomas, 1903, Proc. Zool. Soc. Lond. 2: 201. 

Type locality : Avera, Aroa River, British New Guinea. 
Peroryctes longicauda ornatus (Thomas), Tate, 1948, Bull. Amer. Mus. Nat. Hist. 92: 329. 

Fourteen specimens all from NE. New Guinea: ^50.1121, 1122, $1123, Tapu, 
Upper Ramu River Plateau; $ 1124, Baiyanka, SE. Bismarck Range; $ 1414, 1418, 
$ 1415, 1416, 1417, Kuraka, Kratke Mts. and Kambaidam, Kratke Mts. ; <J 1312, 
$ 1413, Saiko, Bubu River ; $ 1840, 1841, Degabaga, 8 miles east of Hagen Range, Sepik- 
Wahgi Divide ; $ 1842, Menebe, 8 miles east of Hagen Range, Sepik-Wahgi Divide. 

These specimens agree closely with the type except that the five specimens from 
the Kratke Mts. are more rufous, especially ventrally. 

Measurements in mm. (taken in the flesh) : 



us 





-0 

S 




"o 

•8. 

3 









g 

<» « 


IS 


"5) 
1 


a 5 


a S 

11 


£ 

«, 


„ 


N 


50.1121 


3 


260 


196 


59 


25 


6o-o 


23-1 


12-8 


24-8x4-5 


35-9 


5-2 


8-6 


13-2 


9-4 


6-5 


1122 


3 


258 


178 


56 


25-5 


58-6 


22-5 


i3'5 


23-3x4-6 


35-7 


5-2 


8-9 


13-8 


10-3 


70 


1123 


? 


241 


184 


55 


25 


55-1 


22-6 


13-3 


23-3x4-6 


32-9 


4-8 


7-3 


12-6 


9-2 


6-4 


1124 


3 


267 


194 


59 


26 


61-2 


— 


13-2 


26-ox 5-7 


36-7 


5-i 


8-8 


13-1 


9-2 


6-3 


1414 


3 


258 


204 


60 


26 


58-7 


2 3 -0 


13-9 


24-5X5-5 


34-9 


5-8 


7-8 


13-6 


10-5 


7-o 


I418 


3 


257 


207 


56 


26 


6o-6 


23-3 


13-1 


24-8x4-7 


36-1 


5-6 


9-4 


13-7 


100 


6-9 


1415 


? 


258 


187 


54 


24 


58-3 


22-4 


126 


23-9x5-0 


34-2 


— 


8-3 


13-3 


9-7 


6-6 


I416 


9 


239 


185 


56 


27 


58-3 


22-5 • 


12-6 


23-4x4-6 


34-3 


— 


8-5 


13-1 


95 


6-5 


1417 


? 


263 


190 


58 


26 


59-o 


22-6 


12-3 


25-1x4-9 


35-o 


4-8 


8-7 


12-7 


9-3 


6-3 


1412 


3 


275 


217 


61 


26-5 


6i-5 


22-8 


127 


26-8x 5-1 


37-0 


60 


7-6 


14-2 


10-7 


7-3 


1413 


? 


262 


194 


56 


26-5 


57-3 


21-6 


12-0 


25-2x4-5 


33-8 


5-3 


6-9 


12-9 


9-8 


6-8 


1840 





282 


216 


61 


26-5 


62-7 


23-8 


13-8 


27-ox 4-9 


37-3 


5*0 


7-9 


13-6 


io-o 


6-9 


184I 


3 


266 


188 


56 


25 


59-8 


23-4 


136 


25"6x 5-0 


35-5 


5-5 


8-i 


13-6 


9-9 


6-9 


1842 


$ 


265 


196 


54 


25 


597 


22-3 


130 


26-3 x 4-6 


35-9 


4-i 


8-5 


13-4 


9-8 


6-8 



Peroryctes longicauda magna subsp. n. 

Type locality: Ikara, NE. slopes Mt. Simpson, eastern Papua, SE. New Guinea. 

3,500 ft. 
Type: Adult <J 50.1126, collector's No. 768, 18 August 1940. Skin and skull. 
Paratypes: $ 50,1125, collector's No. 751, Ikara, NE. slopes Mt. Simpson, eastern 

Papua, 3,500 ft.; $1128, collector's No. 830, juv. $1129, collector's No. 818, 

juv. (J 1127, collector's No. 884, Enaena, NE. slopes Mt. Simpson, eastern Papua, 

4,000 ft. 

The colour and marking of the skins of this series is almost identical with that of 
the type of Peroryctes longicauda ornata, but the specimens are larger and have 
slightly longer tails, and the undersides are a little darker buff. The general body 
colour is pale brown speckled with black, with a prominent black mid-dorsal line. 
This line begins between the eyes, broadens out on the crown and nape, and continues 
to the base of the tail. There is also a black streak running through each eye from 
the root of the whiskers to the base of the ear, and a black line on each side of the 
rump parallel with the median line, which passes on to the back of the hind legs. 
Unlike ornata there are very few longer hairs on the underside of the tail, but a line 
of longer hairs is usually present along each side. 






IN NEW GUINEA, 1932-1949 291 

The skull is very similar to that of Peroryctes longicauda ornata but is a little 
larger; the additional pair of anterior palatal foramina are smaller and may be 
minute. 

Measurements in mm. of the type and paratypes (taken in the flesh) : 

























g 


3 












■f 1 

■5 




„ 




1 







31 


00 




1. 








1 

§ 


8 

LO 




2 


-s. 

s 
3 


1 


ft 

e 

O .3 


is 

jo § 


"a 


5 


1 






"fe, 


I 


1 


50.1126 Type 


(J 


302 


258 


69 


28 


68-5 


25-1 


29-4x5-6 


13-2 


39-6 


6-i 


8-5 


i5-i 


10-5 


7'4 


1125 


<J 


290 


243 


6 7 


28 


657 


24-0 


28-ox 5-6 


12-9 


39-2 


6-o 


8-5 


i5-i 


II-2 


7-6 


1128 


$ 


303 


226 


63 


27 


76-3 


25-1 


28-1 x 5-2 


13-3 


39-6 


5-5 


8-9 


13-9 


IO-2 


6-8 


1129 


$* 


226 


193 


53 


25 


5i-7 


19-8 


21-8x4-5 


n-8 


31-4 


5-6 


— 


— 


— 


— 


1127 


<?* 


276 


240 


64 


28 


61-7 


23-1 


28-0x5-4 


13-0 


36-9 


6-2 


7-9 


14-9 


io-8 


7-4 



* juvenile 

Peroryctes papuensis sp. n. 

Type Locality: Boneno, Mt. Mura (30 miles NW. Mt. Simpson) Main Range, 
eastern Papua, SE. New Guinea, 4,000-5,000 ft. 

Type: Adult <J 50.1130, collector's No. 816, 3 September 1940. Skin and skull. 

Paratypes: 950.1135, collector's No. 982, 1136, collector's No. 994, Boneno, Mt. 
Mura, eastern Papua, 4,000-5,000 ft. ; $ 1133, collector's No. 865, 1132, col- 
lector's No. 862, 1131, collector's No. 812, juv. 1134, collector's No. 813, Enaena, 
NE. slopes Mt. Simpson, eastern Papua, 4,000-5,000 ft. 

These specimens have the same marking as Peroryctes longicauda ornata, but they 
are much smaller and the general colour of their skins is darker. There is a prominent 
black mid-dorsal line which begins between the eyes, broadens out on the crown 
and nape, and continues to the base of the tail. A black streak runs through each 
eye and there are two short black lines on the rump, one on each side of the middle 
line, which pass on to the back of the hind legs. The pelage on the underside of the 
body is quite a rich orange-buff except in the juvenile specimens where it is light 
grey. As in P. longicauda ornata the hairs on the underside of the tail are longer than 
those on the upper side. 

The skulls are similar to those of P. longicauda ornata, but are much smaller and 
not so heavily built. 

Measurements in mm. of the type and paratypes (taken in the flesh) : 









IS 














3j 


! 


I 


1 








1 

1 














IS 


"« 


.0 


3* 


£ 8 


•2.s 












§ 




s 


K 




bo « 








■a <s 










1 


3 
to 


&S 


£ 


8! 


4 


s 
^ 


$j| 


fe; 




i 


-*«£, 


54 


<\ 


s 


1 


50.1130 Type 


3 


198 


155 


45 


27 


48-5 


17-7 


20-5x4-0 


II-O 


29-6 


4-0 


7-1 


105 


7-4 


5-o 


1X35 


¥ 


196 


150 


45 


28 


48-0 


176 


20-4 X 4-0 


10-7 


28-0 


4-6 


7-7 


9-8 


7-2 


4-7 


1136 


V 


175 


142 


43 


25 


43-9 


16-9 


18-7x3-3 


io-i 


26-2 


4-5 


6-3 


100 


7-5 


4-9 


ii33 


V 


200 


155 


45 


26 


47-4 


17-2 


20-4 x 3-2 


io*6 


28-2 


4-2 


5-6 


9-7 


6-8 


4-7 


1132 


V 


193 


155 


44 


27 


47-4 


17-6 


19-9x3-5 


10-5 


28-6 


4-4 


5-9 


10-2 


7-5 


4-9 


1131 


V 


191 


143 


43 


26 


— 


17-0 


19-9x3-1 


10-4 


27-5 


4-5 


6-o 


9-8 


6-9 


4-6 


ii34 


V* 


127 


105 


32 


21 


33-i 


140 


12-8x3-3 


8-5 


19-4 


3-7 


2-9 


— 


— 


5-0 



juvenile 



292 



MAMMALS COLLECTED BY MR. SHAW MAYER 



Satanellus albopunctatus (Schlegel) 

Dasyurus albopuntatus Schlegel, 1880, (January) Notes Leyden Mus. 2: 51-53. 

Type locality: Arfak Mts., Dutch New Guinea. 
Dasyurus fuscus Milne-Edwards, 1880, (June) Ann. Mag. Nat. Hist. 6: 172. 

Type locality : Arfak Mts. ; Dutch New Guinea. 
Dasyurus daemonellus Thomas, 1904, Ann. Mag. Nat. Hist. 14: 402. 

Type locality: Avera, Aroa River, S. coast, Papua. 
Satanellus albopunctatus (Schlegel), Tate, 1947, Bull. Amer. Mus. Nat. Hist. 88: 142- 



143- 



Fourteen specimens. Thirteen from NE. New Guinea: $50.1393, $1394, juv. 
$ 1395, Buntibasa district, Kratke Mts. ; $ 1392, Kambaidam, Kratke Mts. ; $ 1397, 
1398 (skull and piece of skin), Kuraka, Kratke Mts.; $ 1396, Arau, Kratke Mts.; 
(J 1399, Saiko, Bubu River; $ 1090, 1091, Baiyanka, SE. Bismarck Range; $ 1093, 
$ 1094, Tapu, Upper Ramu River Plateau ; $ 1810, Yanka, eastern slopes Hagen 
Range ; and one $ 1092 from Enaena, NE. slopes Mt. Simpson, eastern Papua. 

This useful series supports the view expressed by Tate (1947) that the three forms 
synonymized above are alike, the seeming differences being mainly due to age. 

Measurements in mm. (taken in the flesh) : 



1 


8 

00 


f 

'si 

e 

&3 


1 



1 

3 




1 


11 

N .0 


-S 
£* 

•IS « 
s £ 


1 


I 


?! 


1 


I 

"a 


S 


50.1094 
1093 

1091 
1090 
1092 
1397 
1398 
1394 
1393 
1392 
1396 
1399 
1810 


9 

0" 

3 
0" 
$ 
<J 
6" 
? 
c? 
6" 
3 
<5 
6" 


241 
231 
262 
250 
275 
271 
283 
255 
283 
279 
280 
298 
269 


221 
224 
244 
247 
280 
270 
259 
253 
271 
277 
264 
290 
239 


43 
46 

44-5 

47 

51 

45 

47 

46-5 

50 

5o 

5i 

54 

46 


29 

27 

27 

29 

27 

29 

30 

29 

3i-9 

29 

3i-5 

29 

30 


50-2 
48-0 
54-3 
53-7 
60-5 

57-8 
52-3 
58-0 
59-3 
59-8 
62-8 
55-6 


34-1 
30-5 
37-6 

37-0 
35-1 
377 
34-7 
39-5 
38-5 
40-1 
397 


12-9 

II-O 

13-6 

I2-I 

14-1 

13-5 

I5'2 

13-6 
15-5 
153 
13-6 
15-0 
9-9 


18-6x7-8 
i8-2X 7-7 
2i-7x 9-6 
22-ox 8-3 
23-0x9-7 

22*7X II-I 
24'7X n-2 

19-9 x 6-7 

23-9x9-8 

25-3X n-8 

22-7x8-8 

25-7x9-5 

2I-OX 8-i 


28-5 
27-4 

30-9 
31-4 

33-8 
29-9 
3i-3 
29-8 
32-7 
32-i 

32-9 

34'9 
30-0 


3-5 
3-7 
37 
4-0 
4-4 

3-6 
2-9 
4-2 
37 
3-0 
5-o 
4-4 


n-8 
12-5 
n-6 
12-8 
12-5 
n-6 
12-3 
12-3 
12-4 
12-3 
12-3 

13-5 

n-5 


7-7 
8-4 
7-8 
8-5 
8-4 
7-7 
8-3 

8-2 

8-3 

8-2 

8-3 
90 
7-5 


4-3 
50 
4-7 
5-o 
4-8 
3-9 
47 
4-7 
47 
4-6 
4-9 
5-2 
4-9 



Neophascogale lorentzi (Jentink) 

Phascogale lorentzii Jentink, 191 1, Notes Leyden Mus. 33: 234. 

Type locality: Hellwig Mts., Dutch New Guinea, 2,600 metres. 
Phascogale nouhuysii Jentink, 191 1, ibid. 33: 235. 

Type locality: Bivak Island, Dutch New Guinea, ± 1,050 metres. 
Phascogale lorentzii venusta Thomas, 192 1, Ann. Mag. Nat. Hist. 8: 358. 

Type locality: Weyland Mts., Dutch New Guinea, 6,000 ft. 

Phascogale venusta rubrata Thomas, 1922, Nova Guinea onder hiding van A. F. Herder schee 

13: 739. 

Type locality: Mount Goliath, central Dutch New Guinea. 
Neophascogale lorentzii (Jentink), Tate, 1947, Bull. Amer. Mus. Nat. Hist. 88: 136. 

Three specimens from Yanka, eastern slopes Hagen Range, NE. New Guinea, 
$ 50.1804, 1805, young ad. $ 1806. 

These are very similar to the specimens from Dutch New Guinea in this Museum, 
but are not so rufous. 



IN NEW GUINEA, 1932-1949 

Measurements in mm. (taken in the flesh) : 



293 







$ 
















^ 


4? 


















.0 








s 








50 


« 




















.^ 




ji 


u 






§ 


*V, 












Ik 

1 








^ 
•« 






§5 


"« 




4s 


.§■1 














to 








v. 




to « 




















i 


£ 


£ 


15 




O s 
^ 




*Z 


2 * 
N 


* 


^ 


I 


^ 


s 


£ 


5 


50.1804 


6" 


171 


188 


40 


23 


44-3 


22-5 


I8-3X6-3 


10-5 


25-4 


4*2 


8-4 


i-o 


3-ox 1-9 


2-7x2-6 


2-7x2-9 


1805 


<J 


200 


213 


42 


24 


5i-o 


26-3 


23-ox 7"8 


10-5 


28-8 


4'9 


8-3 


1*4 


2*9 X 2-0 


2-8X2-5 


2-6x 2-8 


1806 


y. ad. $ 


184 


207 


39 


24 


47-0 


237 


19-5 X 6-o 


io-i 


26-0 


4-5 


8-o 


— 


2-7x1-9 


2-7x2-5 


2-6 x 2-9 



Murexia longicaudata longicaudata (Schlegel) 

Phascogale longicaudata Schlegel, 1866, Ned. Tijdschr. Dierk., Amsterdam, 3: 356. 

Type locality: Aru Islands. 
Murexia I. longicaudata (Schlegel), Tate, 1947, Bull. Amer. Mus. Nat. Hist. 88: 117. 

Seven specimens all from the Kratke Mts., NE. New Guinea: (£50.1400, 1401, 
1402 (skull only), $1403 (skull and piece of skin), Kambaidam; (£1406, Kuraka; 
$ 1405, 1404, Buntibasa district. 

These specimens agree closely with the descriptions of longicaudata particularly 
when comparison is made of the measurements of the molar teeth, which in 
some cases are almost exactly the same as those for the type. They are new to our 
collection. 

Measurements in mm. (taken in the flesh) : 







£ 
















s 



















8 


§ 






^3 




■Vi 




*o 


^ 






•Q 


s 








^2 




3 


^s 


-i2 






i 


CO 






8 


5v 


1 5P 

^ 


N £ 


Co 


Is ^ 


^ 1 


50.1405 


6* 


143 


166 


29 


19-5 


38-5 


22 - 


I5-OX5-0 


7-9 


14-6 


1404 


$ 


130 


160 


28-5 


20 


35-7 


19-5 


I3-8X5-0 


7-8 


14-6 


1403 


¥ 


126 


155 


26-5 


19 


33-3 


187 


11-7X4-8 


7-4 


13-9 


1400 


6* 


140 


155 


28 


20 


37-9 


— 


I3-9X4-9 


7-5 


14-0 


1406 


6* 


150 


182 


30 


21-5 


39-6 


24-0 


I5'9X5-3 


7-4 


14-6 


1402 


— 


— 


— 


— 


— 


41-0 


24-0 


I6-OX5-9 


7-4 


14-6 



1 


►si 


•is 

^ 5 


^ IP 


to 

51b 


Outer corners of 
m 3 


« 










50.1405 
1404 

H03 
1400 
1406 
1402 


21-7 

20-1 

18-5 
20-9 

22-3 
22'7 


35 
3-i 

3-2 

3-7 
3-9 


4'4 
4-5 
4-0 

4-3 
4-6 
4-6 


5-8 
5-4 
5-i 

6-4 
6-6 


12-0 

n-9 
n-7 

13-3 
13-0 


7-6 
7-7 
7-5 
8-o 
7-8 
7-8 


2-7 x 1-9 
2-7 x i-8 
2-6X1-8 
2-7x1-8 
2-7 x 1-9 
2-7x2-0 


2-6x2-3 

2-6 X 2-3 
2-5X2-3 
2-7X2-4 
2-6X2-5 
2-7X2-4 


2-2 X 2'7 
2-3X2-7 
2-2X2-6 
2-3X2-7 
2-3X2-8 
2-3X2-8 


2-OX 2-6 
2-OX2-8 

1-8x2-7 

2-OX 2-9 
2-1 X2-9 
2-OX 2-6 



ZOO. 



Rr 



294 



MAMMALS COLLECTED BY MR. SHAW MAYER 



Murexia longicaudata parva subsp. n. 

Type locality: Baiyanka, Ramu River Divide, SE. Bismarck Range, 7,500 ft. 

Type: Adult <J 50.1114, collector's No. 685, 6 June, 1940. Skin and skull. 

Paratypes: ^50.1117, collector's No. 595, 1118, collector's No. 598, $1119, col- 
lector's No. 593, Tapu, Upper Ramu Plateau, 6,000 ft. ; <$ 1113, collector's No. 
667, 1115, collector's No. 690, 1120, collector's No. 663 [in spirit], $ 1116, 
collector's No. 635, Baiyanka, Purari-Ramu Divide, SE. Bismarck Range, 
7,500 ft. 

This is a small and rather slender race of Murexia longicaudata. The general colour 
of the pelage is similar to that of /. longicaudata but is a little longer and softer, the 
hairs on the back being about 7 mm. long, finely grizzled mouse-grey dorsally and 
silvery grey or buffy grey ventrally ; the hands and feet are pale brown and the long 
tail is covered with short, light brown hairs except at the tip, where for a distance of 
from 10 to 47 mm. or so the hairs are white. The hairs are longer on the underpart 
of the tail and project beyond the tip. 

Skull similar to that of /. longicaudata but much smaller. 

Measurements in mm. of the type and paratypes (taken in the flesh) : 







s 








« 







































^ 




« 

A 


<o 




« 


8 






I 

t3 




t3 




O 


Is 

^ 


*i2 


1- 




to 


<*> 


<S 




$ 


^ 


S ft>o 


do « 




•S * 




fc 


UJ 


S 


s 




s 
^ 


N .5 


£ 


2 s 

^1 -o 


s 




50. 1 1 14 Type 


(J 


132 


175 


26 


19-5 


34-3 


1 9-1 


13-2x4-9 


7-7 


10-9 


1113 


S 


131 


173 


26 


18 


33-3 


l8-2 


11-9x4-1 


7-8 


io-o 


1115 


eJ 


127 


154 


25-5 


20 


32-3 


17-1 


12-4x4-4 


8-o 


9-9 


1116 


¥ 


115 


I48 


23 


17 


30-9 


16-5 


11-0x3-5 


7'3 


9-8 


1119 


¥ 


122 


153 


24 


17 


3i-5 


16-7 


H-7X4-5 


7-4 


9-9 


1117 


<J 


IO9 


I50 


25 


17 


30-0 


— 


10-8x3-7 


7-0 


9-9 


1118 


s 


123 


I6l 


25 


19 


33-o 


17-4 


12-4x3-8 


7-8 


10-4 



1 






1 


41 

.8 I 


"> 
«» si 


3 
s 


Ts 


d 
§ 


"§ 


S 


50. 1 1 14 Type 
1113 

1115 
1116 
1119 
1117 
1118 


6-8 
6-9 
6-8 
6-7 
6-6 
6-9 
7-0 


13-5 
13-0 
13-0 
12-5 
12-4 

12-0 

13-1 


18-6 
18-3 
18-2 
17-0 
17-1 
i6- 5 
18-3 


3-6 

3-4 
3-4 
3H 
3'2 
3-6 


4-0 
3'6 
3-2 
3-6 
3-5 
3-2 
3-6 


5-o 
4.4 

4-5 
4-0 

4-5 
3-9 
4-8 


2-5 X 1-7 
2-5x1-8 
2-5x1-7 
2-4 x i-6 
2-4X i-6 
2-4x1-6 
2-5x1-7 


2-4x2-0 
2-3x2-1 
2-4x2-1 
2-3x2-1 

2*3 X 2-1 
2-3 X 2-0 
2-4X2-I 


2-1 X2'3 
2-1 X2'4 
2-2X2-4 
2-1 X2-3 
2-1 X2'3 
2-OX 2-3 
2-1 X2-3 


1-8x2-4 
1-8x2-4 

I-6X2-I 

1-7x2-4 
1-7x2-2 
1-7x2-3 
1-8x2-4 



IN NEW GUINEA, 1932-1949 



295 



Murexia rothschildi (Tate) 

Phascogale {Murexia) rothschildi Tate, 1938, Novit. Zool. 41: 58. 

Type locality: Aroa River, Papua, probable altitude ± 4,000 ft. 
Murexia rothschildi (Tate), 1947, Bull. Amer. Mus. Nat. Hist. 88: 118. 

Six specimens all from eastern Papua: $ 50.1107, Ikara, NE. slope Mt. Simpson; 
S 1109, 1108, juv. $ 1110 (skull and unstuffed skin), Enaena, NE. slope Mt. Simpson ; 
$ mi, $ 1112, Boneno, Mt. Mura. 

The only specimens of this interesting species which appear to have been previously 
recorded are the type — Tring Museum, Field No. 1, a male, and another male 
collected by the same collector on the same day, A.M.N.H. No. 108106. It is easily 
distinguished from longicaudata and its various races by the broad black mid-dorsal 
stripe. 

Measurements in mm. (taken in the flesh) : 






















"e 


<o 






<0 




3 






-0 

^ 


la 

^ 








so 

It* 


« 


50.1107 


6* 


156 


178 


29 


21-5 


39'8 


20-0 


14-8x6-0 


7-4 


ii-9 


7-7 


1 109 


6* 


154 


184 


27 


20 


38-0 


21-7 


13-0x5-5 


7-2 


n-9 


7-5 


1108 


6* 


I50 


162 


26 


20 


38-i 


21-5 


I3-7X5-5 


7-8 


12-4 


7-8 


IIII 


6* 


132 


163 


27 


19 


34 >0 


i8-8 


12-0x4-0 


7-8 


u-6 


7-8 


III2 


V 


124 


152 


25 


19 


3i-9 


17-8 


11-0x4-0 


7-6 


II-2 


7-7 



Antechinus melanurus (Thomas) 

Phascogale melanura Thomas, 1899, Ann. Mus. Stor. nat. Genova, 40: 191, 
Type locality: Moroka, British New Guinea, 1,300m. 

Phascogale melanura modesta Thomas, 1912, Ann. Mag. Nat. Hist. 9: 9 2 - 
Type locality : Mt. Goliath, Dutch New Guinea. 

Antechinus melanurus (Thomas), Tate, 1947, Bull. Amer. Mus. Nat. Hist. 



: 129. 



Thirteen specimens. Five from eastern Papua: ^50.1103, young ad. ^1102, 
$1104, 1105, Enaena, NE. slopes Mt. Simpson; ^1106, Boneno, Mt. Mura; and 
eight from NE. New Guinea: <} 1100, 1101, Baiyanka, SE. Bismarck Range; $ 1834, 
1835, 1836, (<J 1837, ? x 838, in spirit), Tomba, SW. slopes Hagen Range; $ 1839, 
Degabaga, 8 miles east of Hagen Range, Sepik-Wahgi Divide. 

The general colour of all the specimens is very similar ; the orange patch of hair 
behind the ears is not so well developed in the two specimens from the Bismarck 
Range and the one from Degabaga. 



29 6 MAMMALS COLLECTED BY MR. SHAW MAYER 

Measurements in mm. (taken in the flesh) : 





CO 




S 









^ 

ISO « 


Nasals 


3f 







50.1103 


<* 


123 


142 


23 


17 


31-2 


17-9 


10-9x4-8 


7-1 


9-7 


5'6 


1 102 


y. ad. <J 


in 


137 


22 


17 


29-5 


16-7 


9-8x3-9 


7'3 


9-6 


6-1 


1 104 


? 


115 


143 


22 


17 


30-0 


17-5 


11-2x4-0 


7'3 


9-5 


6-o 


1 105 


? 


112 


135 


21-5 


16-5 


29-8 


17-7 


10-8x4-0 


7-5 


9-9 


6-o 


1106 


s 


112 


130 


22 


16 


28-4 


17-5 


9-8x4-1 


7-2 


9-7 


5-8 


1 100 


<* 


114 


140 


22 


16 


29-0 


17-0 


10-8x4-0 


7'3 


9-7 


6-o 


IIOI 


<J 


103 


122 


21 


16 


26-7 


16-2 


9-8x3-9 


7-2 


9-0 


5-7 


1834 


s 


107 


129 


21 


16 


28-4 


16-3 


10-0x4-5 


7-0 


8-8 


6-o 


1835 


s 


99 


125 


21 


15 


27-0 


15-8 


io-ox 4-2 


6-5 


9-2 


5'9 


1836 


3 


no 


138 


22-5 


15 


30-5 


17-2 


12-0x4-9 


7-2 


9-5 


6-o 


1839 


3 


115 


144 


22 


16 


29-0 


17-5 


11-0x4-0 


7-9 


9-5 


5-7 


1837 


<J 


— 


— 





— 


28-4 


16-5 


io-8 X4*4 


6- 9 


9-2 


6-o 


1838 


$ 


— 


— 





— 


27*9 


16-4 


10-5x4-4 


7-0 


9-1 


5-8 



Antechinus hageni sp. n. 

Type locality: Tomba, SW slopes Hagen Range, Central Highlands, NE. New 
Guinea, 8,200 ft. 

Type: Adult $ 50.1829, collector's No. 1097, 30 June 1947. Skin and skull. 

Paratypes: young ad. $50.1830, collector's No. 1101, (^1831, collector's No. 
mi, 1832, collector's No. 1115, in spirit), Tomba, SW. slopes Hagen Range, 
NE. New Guinea, 8,200 ft. ; $ 1833, collector's No. 1052, Yanka, eastern slopes 
Hagen Range, Central Highlands, NE. New Guinea, 5,500 ft. 

The measurements of this small species are very similar to those for A . wilhelmina 
and from its general body colouring it appears to belong to Tate's A. flavipes group 
which, in New Guinea, contains melanurus, mayeri, centralis, tafa, misim, and 
wilhelmina. The general body colour is a uniform brownish-grey, the bases of the 
hairs grey, the tips yellowish-brown. The rump is not contrastingly reddish as in 
wilhelmina. There are no ear patches. The hairs on the underparts are grey based 
and tipped with white. The hands and feet are pale brown, the digits buffy. The 
tail is brown above and pale buffy below. 

Measurements in mm. of the type and paratypes (taken in the flesh) : 

























"s 










5 






1 
-0 








3 


•U 




3i 


8 


■ft, 










*i 


h. 


















■*» 


"*» 


I 








§ 





"1 




13 


^ 


s 


v 


S* 


-« 

60 « 


<5 


-« 




S '2 
•3 8 


"g 


„ 


„ 


.« 
^ 


k 
v 


£ 


<~n 


£ 


£ 


£3 




8 
O .3 




1 




« 


$t 


T 


6-3 


§ 


£ 


O 


50.1829 Type 


«J 


109 


125 


21 


17-5 


29-6 


16-0 


12-5X4-5 


7-8 


l6"3 


4-3 


15-5 


5-5 


i-4 


89 


1830 


y. ad. $ 


96 


131 


20 


18 


28-3 


15-0 


II-5X3-5 


7-6 


15-2 


4-8 


14-9 


6-3 


5-4 


1-4 


9-0 


1833 


? 


105 


119 


20 


17 


28-2 


14-7 


II-5X3-8 


7-6 


15-5 


3-8 


14-9 


6-3 


5-5 


1-4 


90 


1831 


6" 


— 


— 


— 


— 


29-9 


17-1 


II-5X4-9 


8-o 


16-5 


50 


15-5 


6-3 


5-6 


i-4 


9-4 


1832 


6* 


— 


— 





— 


27-5 


15-0 


— X3-5 


7-8 


— 


C-4-5 


14-3 


6-2 


5-5 


1-4 


8-9 



IN NEW GUINEA, 1932-1949 

Phascolosorex dorsalis whartoni (Tate & Archbold) 



297 



Phascogale [Phascolosorex) dorsalis whartoni Tate & Archbold, 1936, Amer. Mus. Novit., No. 

823:4. 
Type locality: Eastern slope of Mt. Tafa, Central Division of Papua, 2,070 metres. 
Phascolosorex dorsalis whartoni (Tate & Archbold), Tate, 1947, Bull. Amer. Mus. Nat. Hist. 

88: 138. 

Six specimens, all from NE. New Guinea: ^50.1098, 1095, 1097, 1096, $1099, 
Baiyanka, SE. Bismarck Range ; $ 1807, Menebe, 8 miles east of Hagen Range, 
Sepik-Wahgi Divide. 

Measurements in mm. (taken in the flesh) : 







-0 








00 






"e 




^ 




^3 




"O 




-0 


_<o 




-S 


-3 






« 




^ 






5 rSJ 




^ 


■g *« 














^ 










s 




t3 




"« 




O ^ 


"S 


K "e 


"5 ^ 


c/3 






s 

5 






N ,0 


« 
* 






50.1098 


s 


166 


143 


25-5 


21 


39-o 


22-5 


15-7x6-1 


8-3 


i6-i 


1095 


6* 


162 


137* 


26-5 


21 


40-4 


21-2 


I5-7X5-7 


7-0 


16-6 


1097 


6* 


144 


128 


25-0 


19 


37-o 


20-2 


I3-7X5-9 


8-o 


i5-i 


1096 


0* 


138 


135 


25-0 


18 


36-4 


19-8 


13-7x5-6 


8-4 


15-0 


1099 


¥ 


123 


119 


23-5 


18 


34-o 


17-6 


11-8x5-0 


7-9 


14-2 


1807 


? 


119 


123 


24-0 


16 


33'4 


16-5 


12-6x5-4 


8-i 


I3'4 







4! 


5 


Tf 

^ 


§ 







CQ 

g 

to 
O 

S 

O 


n 


5O.IO98 

I095 
IO97 
IO96 
IO99 

1807 


20-8 

22-0 
20-2 
19-4 


3-6 
3-8 
3-6 
3-5 


4-0 
4-0 
4 -i 
4-3 
3-2 


I-O 

i-o 

I-O 

o-8 
0-9 
0-9 


2-5 x i-6 
2-4x1-6 

2-6X 1-7 

2-4X i-6 
2-4 X i-6 
2-4X1-5 


2-3 X 2-0 
2-3X2-0 
2-6X 2-1 
2-3X2-0 
2-3 X 2-0 

2-3x1-9 


2-2 X 2-2 
2-2X2-4 
2-3X2-5 
2-OX 2'3 
2-2 X 2-2 
2-1 X 2-1 


II-I 

IO-7 

II-O 
I I-O 

10-9 
10-3 


7-0 
6-8 
7-5 
6-7 
6-9 
6-8 



Tip broken off. 



Anisomys imitator Thomas 



RODENTIA 



Anisomys imitator Thomas, 1903, Proc. Zool. Soc. Lond. 2: 199-200. 
Type locality: Aroa River, British New Guinea. 

Three specimens, all from NE. New Guinea: $ 50.1159, 1160, Buntibasa district, 
Kratke Mts. ; $ 1161, Saiko, Bubu River. 



298 



MAMMALS COLLECTED BY MR. SHAW MAYER 



Pogonomys macrourus (Milne-Edwards) 

Mus {Pogonomys) macrourus Milne-Edwards, 1877, C.R. Acad. Sci. Paris, 85: 1081. 

Type locality: Arfak, Dutch New Guinea. 
Pogonomys lepidus Thomas, 1897, Ann. Mus. Stor. nat. Genova, 38: 614. 

Type locality: Haveri, Astrolabe Range, Papua. 
Pogonomys lepidus huon Tate & Archbold, 1935, Amer. Mus. Novit., No. 803: 6. 

Type locality: Huon Peninsula, Dutch New Guinea. 
Pogonomys lepidus derimapa Tate & Archbold, 1935, Amer. Mus. Novit., No. 803: 6. 

Type locality: Mount Derimapa, Dutch New Guinea. 

Seventeen specimens. Fourteen from NE. New Guinea: two from the Kratke 
Mts., ^50.1167, Buntibasa district and ^1168 from Kambaidam; six from Dega- 
baga, 8 miles east of Hagen Range, Sepik-Wahgi Divide, <J 1648, 1651, 1650, juv. 
<J 1649, ? J 653, 1652 ; four from Junzaing, Huon Peninsula, $ 1655, 1654, ? 1656, 
1657 > * wo from Mendi, Bismarck Range, $ 1658, $ 1659 '> an d three from eastern 
Papua: juv. ^47.1283, Enaena, NE. slopes Mt. Simpson; juv. ^47.1284, $47.1285, 
Boneno, Mt. Mura. 

The difference in colour between the juvenile and adult pelage is clearly indicated 
in this series. That of the younger animals, especially No. 47.1283, is grey with a 
very light overwash of yellowish-brown ; that of the adult is a bright yellowish-brown. 
The measurements indicate the amount of variability in the species. 

Measurements in mm. (taken in the flesh) : 







* 


-0 














q 


So 


3 
3 
"q 










^ 




^ 




-0 


_ta 




-K» 


£ 
^ 


-^ 






is 

1 




s 





•X3 







1* 

"2 


</5 


"« 


3 
55 


S 






<0 


«3 




S 




^ 


^ 
N^ 


1 




1 




§ 


S 


47-1285 


? 


129 


178 


23 


15 


30-4 


18-9 


II-2 


4'4 


16-4 


4-9 


5-7 


2-5x1-9 


50.1167 


6* 


117 


176 


24 


14 


29-3 


18-0 


io-6 


4'5 


15-5 


4-i 


5-4 


2-4X1-7 


1168 


6* 


125 


175 


24 


15 


30-7 


17-5 


II-I 


4-4 


16-4 


3-3 


5-6 


2-5x1-7 


1648 


0* 


I30 


182 


24 


15 


30-9 


17-5 


u-8 


4-6 


16-3 


37 


5-2 


2-3x1-6 


1651 


0* 


119 


165 


22-5 


14 


29-0 


16-9 


10-7 


47 


15-2 


3-9 


5*2 


2-3x1-7 


1650 


a* 


II 4 


177 


23'5 


13 


29-4 


16-4 


10-9 


4-8 


15-7 


3'9 


5'i 


2-2X1-6 


1653 


? 


127 


I76 


23 


14 


3 o-6 


17-6 


"•3 


4-8 


16-2 


3'9 


4-9 


2-2 X 1-6 


1652 


? 


II 4 


164 


23-5 


14 


28-6 


16-4 


IO'I 


4-6 


150 


3-9 


5-o 


2-3 X 1-6 


1655 


0* 


126 


187 


23 


16 


3 o-8 


17-3 


10-9 


4-9 


16-7 


4'3 


5-5 


2-5X1-7 


1654 


S 


125 


I78 


23 


i6- 5 


31-0 


18-1 


— 


47 


16-5 


4'4 


5-5 


2-5X1-8 


1656 


$ 


129 


173 


23 


15 


30-5 


17-3 


10-5 


4-5 


16-2 


4-5 


57 


2-5X1-8 


1657 


9 


128 


177 


23 


16 


31-2 


18-3 


io-8 


4-i 


16-9 


4-2 


5-4 


2-5x1-7 


1658 


S 


117 


195 


25 


14 


29-8 


17-3 


— 


47 


16-0 


3-8 


5-5 


2-5X1-8 


1659 


? 


115 


2IO 


25 


14-5 


30-8 


i7'5 


10-9 


47 


16-8 


4 -i 


5-5 


2-5x1-8 



Pogonomys mollipilosus (Peters & Doria)* 

Mus mollipilosus Peters & Doria, 1881, Ann. Mus. Stor. nat. Genova, 36: 698. 
Type locality: Katau, Oriomo River, Daru, S. New Guinea. 

Thirty-one specimens. Eight from eastern Papua: ^47.1252, 1253, 1255, juv. 

* See Tate, 1951: 280, for synonyms. 



IN NEW GUINEA, 1932-1949 299 

(J 1251, $47.1244, 1254, I2 56, 1257, Enaena, NE. slopes Mt. Simpson; and twenty- 
three from NE. New Guinea: #47.1242, 1243, Baiyanka, SE. Bismarck Range; 
c2 47.1245, $47.1247, 1249, 1250, juv. $1246, juv. $1248, Tapu, Upper Ramu 
River Plateau; #50.1169, $50.1170, Kambaidam, Kratke Mts. ; #50.1171, 1172, 
$50.1173, 1174, Buntibasa district, Kratke Mts.; #50.1665, 1666, 1667, $50.1668, 
Yandara, Bismarck Range; #50.1661, 1660, $50.1663, 1662, [$50.1661 in spirit], 
Tomba, SW. slopes Hagen Range. 

This series shows a tendency, which is particularly marked in the specimens from 
Yandara and Tomba, for the line of demarcation between the yellowish -brown dorsal 
pelage and white ventral pelage to become indistinct. This is completely so in No. 
50.1660 where the colour of the dorsal pelage merges with that of the under-surface, 
which is yellowish-whitish-grey. 

Pogonomys sylvestris Thomas 

Pogonomys sylvestris Thomas, 1920, Ann. Mag. Nat. Hist. 9: 534. 
Type locality : Rawlinson Mountains, New Guinea. 

Thirty-five specimens. Twenty-three from NE. New Guinea: #47.1268, 1269, 
1273, juv. # 1270a, juv. # 1274, $47.1270, 1270&, Baiyanka, SE. Bismarck Range; 
#50.1162, Saiko, Bubu River; #50.1684, 1685, 1686, $50.1688, 1689, Yandara, 
Bismarck Range; # 50.1693, $ 50.1690, 1691, Yanka, eastern slopes Hagen Range; 
# 50.1695, 1694, $ 50.1696, [# 50.1697, juv. $ 50.1698 in spirit], Tomba, SW. slopes 
Hagen Range; $50.1692, Degabaga, 8 miles east Hagen Range, Sepik-Wahgi 
Divide; and twelve from eastern Papua: # 47.1272a, $47.1271, 1272, Boneno, Mt. 
Mura; #47.1275, 1276, 1277, I2 7^, 1282a, $47.1279, 1280, 1281, 1282, Enaena, NE. 
slopes Mt. Simpson. 

Pogonomys fergussoniensis sp. n. 

Type Locality: Taibutu, Faralulu district, West Fergusson Island. 
Type: Adult # 50.1175, collector's No. 424, 21 July 1935. Skin and skull. 
Paratype: #50.1176, collector's No. 427 (skull only), Taibutu, Faralulu district, 
West Fergusson Island. 

These specimens are most nearly allied to P. mollipilosus. They are larger. The 
general colour is a rusty brown and there is no sharp line of demarcation between 
the dorsal and ventral pelage. Dorsally the pelage consists of grey based hairs with 
russet tips, and longer fuscous hairs. Ventrally the pelage is whitish-buff with 
patches of rust-coloured hairs. The hairs on the fore and hind feet are buff and there 
is a band of dark hairs on the upper side of the wrists. Tail fuscous with light 
patches. Ears fuscous. 

Skull larger and more heavily built than that of mollipilosus; temporal ridges 
prominent. 



3 oo MAMMALS COLLECTED BY MR. SHAW MAYER 

Measurements in mm. of the type and paratype (taken in the flesh) : 




















»5g 

a? 


3f 




"So 


"S 






►o 












O 

4^ 


1| 


-2» 
►if 


■8 






< 
5 






00 


« 




§ 


^ 


§ feo 


tio « 




-JS <S 


e 




"~<» 






s 

^ 


i3 


B 


S3 




S 


N -0 


£ 


is * 

►"-1 -O 


q 


* 


fin 


| 


§ 


50.1175 Type 


6* 


167 


249 


29 


15 


377 


22-8 


14-0 


4'9 


io-6 


20-5 


4'5 


7-3 


3-5x2-4 


1176 


6* 


— 


— 


— 


— 


35-o 


20-7 


12-5 


3'9 


IO-2 


19-3 


4-4 


7*3 


3-5X2-4 



Pogonomys forbesi (Thomas) 

Chiruromys forbesi Thomas, 1888, Proc. Zool. Soc. Lond. 1888: 239. 

Type locality: Sogere, SE. New Guinea. 
Chiruromys pulcher Thomas, 1895, Novit. Zool. 2: 164. 

Type locality: Fergusson Island, D'Entrecasteaux group. (Status fide Rummler.) 
Pogonomys forbesi vultumus Thomas, 1920, Ann. Mag. Nat. Hist. 9: 535. 

Type locality: Milne Bay, SE. Papua. 
Pogonomys forbesi mambatus Thomas, 1920, Ann. Mag. Nat. Hist. 9: 536. 

Type locality: Kokoda, Mambare River, NE. New Guinea. 
Pogonomys {Chiruromys) forbesi satisfactus Tate & Archbold, 1935, Amer. Mus. Novit., No. 
803:7. 

Type locality: Goodenough Island, D'Entrecasteaux group. 
Pogonomys (Chiruromys) pulcher major Tate & Archbold, 1935, Amer. Mus. Novit., No. 803: 8. 

Type locality: Goodenough Island, D'Entrecasteaux group. (Status fide Rummler.) 

Eight specimens. Four from Garaina, Upper Waria River, NE. New Guinea: 
550.1163, 1164, $ 1165, 1166; and four from Wapona, north slope Manaeo Range 
(35 miles NW. Mt. Simpson), eastern Papua: 5 47.1264, $ 1265, 1266, juv. $ 1266a. 



Pogonomys lamia Thomas 

Pogonomys lamia Thomas, 1897, Ann. Mus. Stor. nat. Genova, 38: 615. 

Type locality : Ighibirei, Upper Kemp Welch River, Central British New Guinea. 

Eight specimens, all from eastern Papua: 547.1258, 1259, 1260, Enaena, NE. 
slopes Mt. Simpson; 5 1261, 1267 juv. of $ 1262, 1262a, $ 1262, Boneno, Mt. Mura; 
5 1263, Ikara, NE. ridge Mt. Simpson. 

Pogonomys shawmayeri sp. n. 

Type locality: Taibutu, Faralulu district, West Fergusson Island, 900 ft. 
Type: Adult 5 50.1177, collector's No. 419, 17 July 1935. Skin and skull. 
Paratype: $50.1178, collector's No. 420, Taibutu, Faralulu district, West Fer- 
gusson Island, 900 ft. 

These specimens appear to be closely related to Chiruromys pulcher Thomas, 1895, 
type locality Fergusson Island, which Rummler (1938) describes as a synonym of 
forbesi. They are, however, larger, particularly when compared with the specimens, 
in this collection, of forbesi from the mainland and are at once distinguished by the 



IN NEW GUINEA, 1932-1949 301 

much thicker tail which is covered with coarse brownish-black scales. As in the type 
of pulcher the fur is longer and softer than that of typical forbesi, the hairs in the 
middle of the back being about 15 mm. long. The general colour, however, is very 
similar to that of forbesi, a soft rufous brown instead of the reddish colour of pulcher, 
and the underside is creamy white instead of russet. There is a dark band running 
from the sides of the muzzle which joins the dark ring round the eye. The feet and 
hands are whitish. The tail has coarse scales which are all keeled and almost black 
in colour; in pulcher they are brown and only some are keeled. The skull is 
fairly similar to that of the types of pulcher and forbesi, the most noticeable difference 
being the somewhat broader brain-case (mastoidal breadth 16-2 mm. and c. 16-0 
mm. in shawmayeri; 15-3 mm. in type of pulcher; 14-0 mm. in type of forbesi). 
Measurements in mm. (taken in the flesh) : 



is 

1 


8 



s 











fa 

ts 


3 

^ 
4^ 


« 


1 


8 
g 








£ 


£ 


S 


b? 


€ 


s 
O .3 




8 iS 


1 


Q 


£ 


« 


s 


a 


50.1177 Type 


6* 


156 


248 


29 


20-5 


36-9 


C.22-5 


6-o 


I2-6X 4-0 


II-O 


5-o 


19-0 


5-8 


2-6 x 1-9 


1178 


V 


155 


232 


29 


20-5 


— 


21-7 


57 


n-8x 3-9 


io-8 


4-6 


18-1 


5-7 


2-6x 1-9 



Hyomys goliath goliath (Milne-Edwards) 

Mus goliath Milne-Edwards, 1900, Bull. Mus. Hist. Nat., Paris, 6: 165. 

Type locality: Aroa River, British New Guinea, 3,000-7,000 ft. 
Hyomys meeki Thomas, 1903, Proc. Zool. Soc. Lond. 2: 198. 

Type locality : Avera, Aroa River, British New Guinea. 

Fifteen specimens, all from NE. New Guinea: $50.1179, Kuraka, Kratke Mts. ; #1180, 
1181, juv. # 1183, juv. # 1182, $ 1184, Buntibasa district, Kratke Mts. ; # 1185, 
1186, Arau district, Kratke Mts. ; juv. $ 1187, Saiko, and # 1188, Bubu River 
district ; $ 1189, 1190, Zageheme, Cromwell Mts., Huon Peninsula [in spirit] ; # 1681, 
$ 1682, Yanka, eastern slopes Hagen Range; juv. $ 1683, Menebe, 8 miles east of 
Hagen Range, Sepik-Wahgi Divide. 



Mallomys rothschildi Thomas subsp. 
Range: New Guinea and (?) Flores. 

Forty-eight specimens. Thirteen from eastern Papua: young ad. #47.1344, 
$1345, 1346, 1347, Boneno, Mt. Mura; young ad. $1341, Ikara, NE. slopes Mt. 
Simpson; # 1349, *35o, young ad. #1348, 9 1352, 1353, 1355. young ad. $1354, 
juv. $ 1351, Enaena. NE. ridge Mt. Simpson ; and thirty-five from NE. New Guinea: 
#50.1193, 1194, 1195, juv. #1196, $1197, 1198, Saiko, Bubu River; #1192, 
Buntibasa district, Kratke Mts. ; $1191, Kuraka, Kratke Mts. ; #47.1342, $47.1343, 
Tapu, Upper Ramu River Plateau; #47.1334, 1335, 1339, $47.1336, 1337, juv. 
$ 1338, young ad. $1340, Baiyanka, SE. Bismarck Range; #50.1780, 1781, 1782, 
Guyebi, Bismarck Range; #1784, 1785, 1786, 1787, juv. #1783, $1788, Yanka, 
eastern slopes Hagen Range ; # 1789, 1790, $ 1791, Tomba, SW. slopes Hagen Range ; 



zoo. i, 10 



ss 



302 



MAMMALS COLLECTED BY MR. SHAW MAYER 



<J 1792, $ 1793, I794> 1795 » Menebe, 8 miles east of Hagen Range, Sepik-Wahgi 
Divide ; $ 1796, $ 1797, Degabaga, 8 miles east of Hagen Range, Sepik-Wahgi Divide. 

This excellent collection of skins from a number of localities, ranging from the 
Hagen Range in the north to Mt. Simpson in the south, indicates that there is a 
great deal of variation in the colour and texture of the pelage of this species, from 
blackish and brownish-grey to dark brown. Both grey and brown forms occur 
together though the really dark brown specimens have so far only occurred in collec- 
tions from the Bismarck and Hagen Ranges. Another variation in coat colour, which 
occurs in both grey and brown forms, is the presence of a band of white hairs across 
the middle of the underside which may go round on to the back (Nos. 50.1786, 50.1790 
and 50.1791). Specimen No. 50.1789 has a few white hairs in the middle of its side. 

Measurements in mm. (taken in the flesh) : 



1 




1 

i 


H 
$ 


I 


1 



s 


K 
4 


1 

5j 


la 

N .0 


1 


1 


3* 

* 1 


§ 


Ikara, Mt. Simpson, eastern Papua 
Boneno, Mt. Mura, eastern Papua 

Enaena, Mt. Simpson, eastern Papua 

Saiko, Bubu River, NE. New Guinea 

Buntibasa district, Kratke Mts. NE. 

New Guinea 
Kuraka, NE. New Guinea 
Tapu, Upper Ramu River Plateau, 

NE. New Guinea 

Baiyanka, Bismarck Range, NE. New 
Guinea 

Guyebi, Bismarck Range, NE. New 
Guinea 

Yanka, Hagen Range, NE. New 
Guinea 

Tomba, Hagen Range, NE. New 
Guinea 

Menebe, Nr. Hagen Range, NE. New 
Guinea 

Degabaga, 8 miles east of Hagen 
Range, Sepik-Wahgi Divide, NE. 
New Guinea 


47-1341 
/ 47-1344 
I 1345 
| 1346 

V 1347 
/ 47-1349 

I350 

1348 

{ 1352 

I 1353 

1 1355 

V 1354 
f 50.1193 

1194 

( 1195 
1197 

V 1 198 
50.1192 

50.1191 
i 47.1342 
I 1343 
/ 47-1334* 
1335* 
/ 1339 
\ 1336* 
1 1337* 

V 1340 

I 50.1780* 

1781 

I 1782 
, 50.1784* 
1785* 
j 1786* 
I 1787* 
v 1788 
/ 50.1789* 
{ 1790* 
I 1791 
/ 50.1792* 
1 1793 
1 1794 
1 1795 
1 1796 
j 1797 


y. ad. ? 

y. ad. 6* 
? 
$ 
$ 
0* 
6* 

y. ad. 6* 
? 
$ 
$ 

y. ad. $ 
6* 
0* 

9 
? 
3 

$ 
6* 
$ 

a 
a 
a 

$ 
? 

y. ad. $ 
<J 
6* 
<? 
3 
6* 
6* 

a 

? 

6" 
0* 

$ 

a 
$ 
$ 
? 
a 
$ 


363 

359 
37i 
410 
406 
374 
377 
362 
354 
384 
376 
357 
384 
398 
390 
360 
378 
346 

368 
378 
322 
39i 
370 
375 
357 
376 
374 
367 
360 
376 
384 
364 
353 
365 
406 
355 
369 
344 
350 
372 
401 
385 
389 
363 


380 
396 
382 
400 
416 
368 
391 
408 
386 
387 
416 
386 
382 
360 
400 
365 
384 
416 

375 

350 

335 

395 

382 

295t 

413 

410 

394 

380 

394 

364 

400 

396 

354 

382 

353 

385 

337 

348 

380 

355 

408 

405 

373 

378 


70 

76-5 

68 

74 

74 

69 

72 

74 

69 

73 

72 

7i 

72 

67-5 

72 

7o 

7i 

70 

67 

68 

65 

7o 

66 

74 

70 

70 

73 

68 

73 

70 

7o 

68 

70 

7i 

7i 

7o 

68 

68 

68 

7i 

76 

7i 

71-5 

69 


29 
29 
30 
30 
30 
30 
30 
29 
28 
30 
29 
27 

28-5 

3i 

30 

29 

3i 

28 

27-5 

27 

27 

30 

30 

27 

30 

30 

28 

28 

30 

29-5 

30 

29 

30 

30 

33 

3i 

29 

29 

28 

3i 

29 

27 

27 

26 


72-9 
79-3 

72-6 

75-9 
74-2 

73-9 
70-3 

71-0 
71-9 

71-5 
71-1 

70-1 
69-8 
75-2 


36-4 
41-8 

40-0 

397 
38-9 

37-4 
3 8-i 

36-0 

38-i 

39-4 
37-0 

36-4 
367 
407 


39-6 
43-0 

40-4 

42-4 
437 

41-4 
40-0 

39-8 
40-1 

40-1 
40-5 

39-4 
39-6 
44-1 


297 
32-0 

28-8 

307 
30-4 

29-8 
27-0 

287 
27-4 

28-0 
29*4 

27'9 
28-0 
33-9 


15-2 
15-5 

15-3 

15-5 
15-3 

14-6 
14-1 

13-6 

14-1 

14-0 
15-2 

I3'9 
14-2 
I4'5 


17-0 
17-1 

17-5 

163 

17-4 

177 
17-0 

16-6 
17-1 

17-0 
17-0 

17-5 
17-0 
17-8 



* Dark brown specimens. 



t Tip broken off. 



IN NEW GUINEA, 1932-1949 303 

Rattus exulans exulans (Peale) 

Mus exulans Peale, 1848, U.S. Exploring Expedition. . . . 1838-42. Under the command of 
C. Wilkes, 8: 47, Philadephia. 
Type locality: Fiji Isands. 

Three specimens from Tongoa Island, New Hebrides: ^50.1200, juv. $50.1201 
skull only, ?5o.i202 skull only. 

Rattus exulans browni (Alston) 

Mus browni Alston, 1877, Proc. Zool. Soc. Lond. 1877: 123. 

Type locality: Duke of York Island. 
Mus echimyoides Ramsay, 1877, Proc. Linn. Soc. N.S.W. 2: 15. 

Type locality : Duke of York Island. 

Nineteen specimens. Fourteen from NE. New Guinea: ^50.1205, 1206, Kam- 
baidam, Kratke Mts. ; ^47.1131, 1132, 1133, $47.1134, Tapu, Upper Ramu River 
Plateau; $47,1135, Baiyanka, SE. Bismarck Range; <J 50.1751, 1752, 1753, 1754, 
x 755> $50.1756, 1757, Yandara, Bismarck Range; three from eastern Papua: 
<£ 47.1138, Ikara, NE. slopes Mt. Simpson; # 47.1139, $ 47.1140, Boneno, Mt. Mura; 
and two, ^47.1136, $47.1137, from Lau, Bainings Mts., Gazelle Peninsula, New 
Britain. 

Rattus ruber tramitius Thomas 

Rattus mordax tramitius Thomas, 1922, Ann. Mag. Nat. Hist. 9: 262. 

Type locality: Mamberano-Idenburg region (Doormanpad-bivak), N. Dutch New Guinea. 
Rattus leucopus utakwa Rummler, 1935, Z. Saugertierk. 10: 115. 

Type locality: Camp No. 3, Utakwa River, 2,000 ft. 
Rattus mordax hageni Troughton, 1937, R ec - Aust. Mus. 20: 120. 

Type locality: Upper Wahgi River, south slopes of Mt. Hagen, south of Sepik Division, New 
Guinea. 

Nineteen specimens. Seventeen from NE. New Guinea: three, <$ 50.1208, 1209, 
$1210, from Saiko, Bubu River; twelve, ^1737, 1738, 1740, 1741, 1742, 1739, 
$ 1748, 1746, 1747, 1743, 1745, 1744, from Yandara, Bismarck Range; two, (J 1749, 
$ 1750, from Yanka, eastern slopes Hagen Range ; and two from eastern Papua : 
$47.1156, Enaena, NE. slopes Mt. Simpson; $47.1159, Boneno, Mt. Mura. 

Tate (1951 : 331, 333) suggests that Rattus ruber hageni from the Mt. Hagen area, 
north-east New Guinea, is possibly the same as R. ruber tramitius from the mountains 
south of the Idenburg River, north Netherlands New Guinea, with which utakwa 
from south-west Netherlands New Guinea is synonymized. Two of the nineteen 
specimens in this collection came from the eastern slopes of Mt. Hagen ; twelve from 
the neighbouring Bismarck Range ; three from the Bubu River (Upper Warai River), 
south-eastern north-east New Guinea; one from Mt. Mura; and one from Mt. Simp- 
son, eastern Papua. The general colour of the specimens varies from the 'buffy- 
ochraceous' of the type of hageni to the 'blackish-grey very finely ticked with 
buffy' of the type of tramitius, and the range of the measurements of the skins 
and skulls includes those of tramitius and hageni, so that it is impossible to sepa- 
rate the two. 



3°4 



MAMMALS COLLECTED BY MR. SHAW MAYER 



Tate (195 1 : 333) states that the mammary formula of hageni is unknown and that 
therefore it may be a race of verecundus or leucopus. But Troughton in his descrip- 
tion of the type gives the mammary formula as 2-2=8. 

The following are the measurements of the skins of six adult males and ten 
adult females, and of the skulls of seven adult males and six adult females, of 
tramitius. 











Standard 




Extremes 


A verage 


deviation 




3 


9 


$ 


9 


3 


9 


Head and body .... 


152-175 


135-172 


162 


152 


7-8 


9-2 


Tail . 






133-155 


127-153 


144 


135 


9-2 


7-5 


Hind foot . 






32-35 


3°-34 


34 


32 


i-3 


i-6 


Ear . 






1 9-21-5 


18-21 


19-8 


19 


1-2 


1-2 


Condylo-basal length 






37-39-3 


34-9-39-6 


38-7 


37-2 


I-O 


1-7 


Zygomatic breadth 






18-3-21-4 


18-7-21-2 


19-7 


199 


I-O 


I-O 


Inter-orbital breadth 






5-8-6-1 


5-6-6-2 


5-9 


5-9 


o-i 


0-2 


m 1 ^ . 






6-6-7-3 


6-7-7-4 


7-0 


7-1 


o-3 


o-3 


m 1 length . 






3-2-3-9 


3-1-3-6 


3-5 


3-4 


0-3 


0-2 


m 1 breadth 






2-1-2-6 


2-1-2-6 


2-3 


2-3 


0-2 


0-2 



Rattus ruber fergussoniensis subsp. n. 

Type locality: Faralulu district, West Fergusson Island, SE. New Guinea, c. 900 ft. 
Type: Adult $ 50.1211, collector's No. 436, 31 July 1935. Skin and skull. 
Paratype: $50.1212, collector's No. 441, skull only, Faralulu district, West 
Fergusson Island, SE. New Guinea, c. 900 ft. 

This short-tailed rat is most closely related to R. ringens feliceus (Ellerman, 1949) 
and R. ringens coenorum (? = bandiculus) . Its size, proportionate length of tail to 
body, and the size of the scales on the tail (6-7 rings per cm.) make it very similar 
to feliceus. The general colour, however, is much darker and is similar to that of 
coenorum, a grizzled brownish-grey, only it is suffused with russet. This colour 
occurs in irregular streaks on the sides and under surface which is otherwise buffy 
grey. 

The skull is not quite as large as, but is most closely allied to, that of the type of 
bandiculus, which may be synonymous with coenorum [see Tate, 1951, p. 332). 
The palatal foramina are straighter and narrower and the molar teeth are arranged 
in a slight curve instead of in the straight almost parallel lines of coenorum. 

Measurements in mm. of the type and paratype (taken in the flesh) : 



1 

s 

1 


CO 


f 

S 


1 




s 





1 £ 


.0 

11 
N J 


1 




q 


! 

1 


s 

'I 


"g 


1 


50.121 1 Type 


6* 


225 


194 


41 


22 


50 


25-3 


19-5x6-2 


6-8 


3-4 


28-7 


9-6 


8-9 


4-3X2-5 


1212 


V 


— 


— 


— 


— 


46-9 


25-0 


19-0x5-2 


7-o 


2-3 


27-7 


8-6 


8-6 


4-0X2-5 



IN NEW GUINEA, 1932-1949 305 

Rattus ruber rosalinda Hinton 

Rattus rosalinda Hinton, 1943, Ann. Mag. Nat. Hist. 10: 557. 

Type locality: Tapu, Upper Ramu River Plateau, NE. New Guinea. 

One additional specimen, $ 50.1207 from Kambaidam, Kratke Mts., NE. New 
Guinea, to the eight specimens which include the type from Tapu, NE. New Guinea. 

Rattus niobe haymani Ellerman 

Stenomys klossi Thomas, 191 3, Ann. Mag. Nat. Hist. 12: 207 (preoccupied). 

Type locality: Upper Utakwa River, Dutch New Guinea, 5,500 ft. 
Rattus niobe haymani Ellerman, 1941, The families and genera of living rodents, 2: 206 (new 



One specimen, <$ 50.1765, from Yanka, eastern slopes Hagen Range, Central 
Highlands, NE. New Guinea. 

Rattus verecundus tomba subsp. n. 

Type locality: Tomba, SW. slopes Hagen Range, Central Highlands, NE. New 

Guinea, 8,500 ft. 
Type: Adult <J 50.1766, collector's No. 1093, 27 June 1947. Skin and skull. 

This specimen seems to be most nearly allied to R. v. mollis Rummler, 1935, from 
Morobe, Mt. Misim, Papua, 5,850 ft. It is smaller than R. v. verecundus. The pelage 
is fine, long and soft, and the hairs on the back, which are about 6 mm. long, are 
dark grey tipped with yellowish-brown. On the under surface they are slate-grey 
tipped with white; a few are tipped with yellow. The feet and hands are white 
and there is a white spot on the chest. The tail is covered with short fine yellowish- 
brown hairs except for about 42 mm. at the tip where the hairs are white. 

The skull is smaller and lighter than that of R. v. verecundus ; the temporal ridge 
is barely visible ; and the anterior palatal foramina are pointed at both ends, not 
more rounded posteriorly as in R. v. verecundus. 

Measurements in mm. of the type (taken in the flesh) : 



.0 
3 


to 




IS 

1 


'I 




5 


K 

1 


1 

B 

-0 


1 £ 


a 


Is 

^ 
N .0 


s 

1 




Q 


1 


>g 

£5 


1 


50.1766 Type 


6* 


136 


146 


32 


19 


32-3 


18-4 


i6- 4 


13-3 


5-9 


9-0 


5-6x2-7 


50 


5-8 



Rattus shawmayeri Hinton 

Rattus shawmayeri Hinton, 1943, Ann. Mag. Nat. Hist. 10: 556. 

Type locality: Baiyanka, Purari-Ramu Divide, SE. Bismarck Range, NE. New Guinea. 

Eight specimens all from NE. New Guinea: <$ 50.1763, $ 1764, duplicate collector's 
No. 1 143 (skull and piece of skin), high slopes Mt. Wilhelm, Bismarck Range ; $ 1758, 
Yandara, Bismarck Range; ^1762, Bogo, south slopes Bismarck Range; #1761, 
1760, Tomba, SW. slopes Hagen Range ; $ 1759, Yanka, eastern slopes Hagen 
Range. 



306 



MAMMALS COLLECTED BY MR. SHAW MAYER 



These specimens are a useful addition to our collection in which, so far, the type of 
the species has been the only representative. 
Measurements in mm. (taken in the flesh) : 







* 








'st 




►si 




►« 


3| 










* 








^ 




Is 


I 3 




3 







m 


►« 
►0 


s 


<a 


<5 




cs 


O S 


?s $> 


<s 




« 


§ <o 








£ 


CO 


tcj 


h 


H3 


^ 


O ^ 


N^ 


^ 


Dh 


^< 


Q 


§ 


s 


50-1763 


6* 


103 


158 


23 


18 


26-9 


15-6 


9'9 


4'3 


14-0 


4'6 


7-0 


4*2 


i-3 


1764 


¥ 


99 


159 


23 


16-5 


25-6 


i5'9 


9'9 


4-3 


13-5 


4-6 


7-0 


4'3 


i-3 


1758 


a 


IOI 


168 


24 


17*5 


27-0 


16-1 


10-4 


4'4 


14-6 


5-2 


7-8 


4-0 


1-2 


1762 


a 


100 


I46 


24 


16-5 


26*2 


157 


9-9 


4*2 


13-8 


5-o 


7*3 


4-3 


1-3 


1761 


s 


107 


158 


24 


19 


27-2 


16-3 


9-9 


4*3 


I4'5 


4*5 


7-3 


4'2 


1-3 


1760 


6* 


104 


I50 


24-5 


18 


20-0 


15*3 


9-8 


4'5 


13-8 


47 


7-2 


4-3 


i'3 


1759 


¥ 


114 


155 


25 


19 


27*4 


16-2 


10-3 


4*7 


14-8 


5'4 


8-2 


4-3 


i-3 



Melomys levipes clarae (Rummler) 

Melomys levipes clarae Riimmler, 1935, Z. Sdugertierk. 10: 108. 
Type locality: Sumuri Mountain, Weyland Mountains, 2,000 ft. 

Two specimens both from NE. New Guinea: $50.1715, Degabaga, 8 miles east 
Hagen Range, Sepik-Wahgi Divide ; $ 1716, Menebe, 8 miles east Hagen Range, 
Sepik-Wahgi Divide. 

These extend the range of clarae to NE. New Guinea and to an altitude of 4,500- 
6,000 ft. The type of M. I. weylandi was taken at 5,000 ft. but the measurements of 
the skulls of these specimens agree with those of the type of clarae. 



Melomys levipes subsp. 

Thirteen specimens identified by Ellerman as a subspecies of M. levipes. Eleven 
from NE. New Guinea: $47.1202, 1203, 1204, 1205, 1206, 1207, $ 1208, 1209, 1210, 
1211, 12110, Baiyanka, SE. Bismarck Range; and two from eastern Papua, $ 1212, 
Enaena, Mt. Simpson; ? 1213, Ikara, NE. ridge Mt. Simpson. 
Measurements in mm. (taken in the flesh) : 



J 

§ 

1 




t 

1 

I 


s 


s 


SI 


1 



^ 


li 

bo « 
N .0 


J8 

s 

1 


3 
•« 

^1 


5 

"bo 
1 

"« 


31 


M 


% 


47-1202 


6" 


156 


158 


35 


20 


35-4 


17-5 


14-3 


6-4 


19-9 


6-3 


8-o 


3-8x2-3 


1203 


6* 


150 


153 


35-5 


20 


357 


17-4 


14-8 


6-6 


20-0 


6-3 


8-o 


3-8x2-2 


1204 


<? 


157 


160 


35 


20 


36-6 


17-3 


I5-I 


6-6 


20-9 


6-5 


7-3 


3-8x2-3 


1205 


6* 


157 


156 


35 


20 


34-5 


17-9 


13-8 


7-o 


I9-0 


6-o 


7-8 


3-9x2-4 


1206 


<? 


154 


156 


35 


21 


35*7 


17-9 


15-3 


6-9 


20-0 


5-7 


7-9 


3-8x2-3 


1207 


<J 


153 


156 


35 


19 


35-6 


17-8 


14-8 


6-8 


20-4 


6-5 


7-9 


3-9X2-3 


1208 


V 


146 


140 


33-5 


19 


34-2 


17-9 


13-9 


6-6 


19-4 


6-7 


7-8 


3-9x2-3 


1209 


V 


159 


166 


36 


20 


357 


17-7 


14-3 


6-7 


20-0 


6-2 


7-9 


3-9x2-4 


I2IO 


V 


152 


144 


35 


19-5 


35-3 


17-9 


14-5 


6-5 


20-0 


6-i 


7-9 


3-8x2-4 


I2II 


¥ 


150 


153 


35 


21 


34-7 


17-4 


i5-o 


6-6 


19-5 


6-o 


8-i 


4-0 X 2-4 


1211a 


V 


144 


144 


35'5 


19 


35-8 


17-0 


13-8 


6-5 


19-0 


6-o 


7-8 


3-8x2-3 



IN NEW GUINEA, 1932-1949 



307 



Melomys moncktoni moncktoni (Thomas) 

Uromys moncktoni Thomas, 1904, Ann. Mag. Nat. Hist. 14: 399. 
Type locality: NE. New Guinea. 

Six specimens, all from NE. New Guinea: ^50.1709, 1710, $1711, 1712, high 
northern slopes Mt. Wilhelm, Bismarck Range; $ 1713, Yandara, Bismarck Range; 
$ 1714, Yanka, eastern slopes Hagen Range. 

Melomys lutillus lutillus (Thomas) 

Uromys lutillus Thomas, 19 13, Ann. Mag. Nat. Hist. 12: 216. 
Type locality: Owagarra, Angabunga River, Central Division, Papua. 

One specimen, ? 47.1214, from Enaena, NE. slopes Mt. Simpson, eastern Papua. 

Melomys rufescens rufescens (Alston) 

Uromys rufescens Alston, 1877, Proc. Zool. Soc. Lond. 1877: 124. 

Type locality: Duke of York Island, between New Britain and New Ireland. 
Mus musavora Ramsay, 1877, Proc. Linn. Soc. N.S.W. 2: 16. 

Type locality: Duke of York Island. 

Twenty-six specimens. Fourteen from NE. New Guinea: ^47.1193, 1195, 1196, 
juv. <2 1 194, ? 1197, 1198, 1198a, Tapu, Upper Ramu River Plateau; J 47.1199, 
$ 1200, 1201, Baiyanka, SE. Bismarck Range ; $ 50.1213, Kambaidam, Kratke Mts. ; 
$50.1701, Yandara, Bismarck Range ; $ 50.1700, Yanka, eastern slopes Hagen Range ; 
$50.1699, Tomba, SW. slopes Hagen Range; and twelve from eastern Papua: 
(£47.1184, 1185, 1186, 1187, juv. (£1183, $1188, 1189, 1190, 1190a, 11906, 1191, 
1192, Enaena, Mt. Simpson. 

Melomys rufescens dollmani Rummler 

Melomys rufescens dollmani Rummler, 1935., Z. Saugetierk. 10: 106. 
Type locality: Buntibasa district, Kratke Mts., NE. New Guinea. 

Four specimens, all from NE. New Guinea: one ^47.1215, from Tapu, Upper 
Ramu River Plateau, and three $$ 50.1718, 1719, 1717, from Tomba, SW. slopes 
Hagen Range. 

Measurements in mm. (taken in the flesh) : 

























oo 










* 




"0 




-0 


<o 




1 














« 
•« 









** 




^ 




5 


s 

HI 








« 




8 


v 


8 *>o 


b.0 « 




-S « 




"S S 






£ 


<» 

Sq 


« 
^ 


5 


« 

^ 


8 


N -0 


IO-O 


£ * 

»~~i £ 


ft, 


s £ 

<*;«£, 


"s 


§ 


47-1215 


6* 


130 


184 


28 


18 


3i-i 


16-7 


5'7 


1 6-1 


4-6 


5'9 


2-9 x i«9 


50*1718 


¥ 


134 


175 


29 


17-5 


31-6 


17-5 


10-7 


5-5 


i6-6 


4-5 


6-2 


3-ox i-9 


1719 


9 


141 


191 


29 


18 


31-6 


16-5 


1 1 - 4 


5'5 


16.4 


4-2 


6-o 


3-0x1-9 


1717 


9 


144 


156* 


29'5 


18 


32-3 


18-0 


1 1 -5 


5-8 


17-2 


4-6 


6-1 


3-ox i-9 



Tip broken 



3 o8 



MAMMALS COLLECTED BY MR. SHAW MAYER 



Melomys fellow si Hinton 

Melomys fellowsi Hinton, 1943, Ann. Mag. Nat. Hist. 10: 554. 

Type locality: Baiyanka, SE. Bismarck Range, NE. New Guinea, 8,000. 

Seventeen specimens, all from NE. New Guinea ; type <£ 47.1175, paratypes $ 1174, 
1176, 1177, $1178, 1179, 1180, 1181, juv. $ 11810, [? 1182 in spirit], Baiyanka, SE. 
BismarckRange ; $50.1704, Yandara, Bismarck Range ; $50.1705, 1706, high northern 
slopes Mt. Wilhelm, Bismarck Range ; $ 50.1707, [# 1708 in spirit], Tomba, SW. slopes 
Hagen Range; (£50.1702, $ 1703, Yanka, eastern slopes Hagen Range. 

The first ten specimens were mentioned by Hinton in 1943 when describing the 
type. The other specimens are all very similar to these. 



Pogonomelomys sevia tatei Hinton 

Pogonomelomys tatei Hinton, 1943, Ann. Mag. Nat. Hist. 10: 554. 

Type locality: Baiyanka, Purari-Ramu Divide, SE. Bismarck Range, NE. New Guinea, 
8,000 ft. 

Eight adult specimens, all from NE. New Guinea: seven from Tomba, SW. slopes 
Hagen Range, $ 50.1721, 1720, $ 1722, 1723, 1724, [^ 1725, ? 1726 in spirit], and 
one from the high northern slopes of Mt. Wilhelm, Bismarck Range, $ 1727. 

These are a useful addition to our collection which only contained the type and 
two young paratypes. The general colour of all six specimens is a rich reddish-brown. 
This is the colour of the adult pelage ; that of the young specimens is much greyer. 

Measurements in mm. (taken in the flesh) : 



.8 
1 


to 


t 
s 







K 





60 <$ 


"Kb 


2 
? S 


1 


f 


s 

1 

3 






£ 


£ 


£ 


5 


4 


s 




1 




3 


« 


* 


s 


% 


50.1721 


6* 


135 


173 


25 


17-5 


33-2 


19-3 


n-5 


6-o 


9-3 


17-6 


6-2 


6-5 


3-0x17 


1720 


<? 


135 


166 


25 


18 


3i-6 


18-2 


C. 12*0 


5-3 


8-4 


16-5 


5'6 


6-4 


2-9X i-8 


1722 


? 


138 


183 


25 


18 


32-8 


18-2 


117 


5-o 


8-7 


17-3 


5'5 


6-i 


2-8x i-8 


1723 


9 


128 


172 


25 


18 


30-8 


17-9 


9-8 


5-3 


8-5 


i6-o 


5-3 


60 


2-8x 17 


1724 


? 


120 


180 


24 


18 


30-5 


17-7 


ii-i 


4-9 


8-4 


i6-i 


5-4 


6-3 


3-ox i*8 


1727 


9 


127 


184 


24 


17 


30-6 


i8-i 


n-4 


5-9 


8-5 


16-5 


5-5 


6-2 


2-8x 1-7 


1725 


S 


— 


— 


— 


— 


— 


i8-o 


II-O 


5-7 


8-8 


17-0 


5-2 


6-5 


3-0x1-7 


1726 


9 


— 


— 


— 


— 


31*6 


17-3 


ii-i 


5-2 


8-6 


i6-6 


5-8 


60 


2-8x 17 



Uromys anak Thomas 

Uromys anak Thomas, 1907, Ann. Mag. Nat. Hist. 20: 72. 

Type locality: Ifogi, Brown River, NE. Papua, ± 4,000ft. 
Uromys rothschildi Thomas, 191 2, Nov. Zool. 19: 91. 

Type locality: Rawlinson Mts., Huon Peninsula, New Guinea. 

Nine specimens all from NE. New Guinea: <J 50.1227, $ 1228, Buntibasa district, 
Kratke Mts. ; $ 1229, Kuraka, Kratke Mts. ; $ 1232 skull only, Apimuri, Kratke Mts. ; 
c? 1230, 1231, Saiko, Bubu River; ^1676, juv. ^1675, Degabaga, 8 miles east of 
Hagen Range, Sepik-Wahgi Divide ; (J 1677, Menebe, 8 miles east Hagen Range, 
Sepik-Wahgi Divide. 



IN NEW GUINEA, 1932-1949 309 

Uromys caudimaculatus aruensis Gray 

Uromys aruensis Gray, 1873, Ann. Mag. Nat. Hist. 12: 418. 

Type locality: Aru Islands. 
Uromys validus Peters and Doria, 1881, Ann. Mus. Stor. nat. Genova, 36: 703. 

Type locality: Katau, mouth of Fly River, Papua. 
Hapalotis papuanus Ramsay, 1883, Proc. Linn. Soc, N.S.W. 8: 18. 

Type locality: New Guinea. 
Uromys nero Thomas, 1913, Ann. Mag. Nat. Hist. 12: 208. 

Type locality: Camp No. 3, Utakwa River, Dutch New Guinea, 2,500 ft. 
Uromys scaphax Thomas, 1913, Ann. Mag. Nat. Hist. 12: 209. 

Type locality: Canoe Camp, lower Setakwa River, Dutch New Guinea, 150 ft. 
Uromys prolixus Thomas, 1913, Ann. Mag. Nat. Hist. 12: 213. 

Type locality: Haveri, Astrolabe Range, Papua, 2,000 ft. 
Uromys ductor Thomas, 191 3, Ann. Mag. Nat. Hist. 12: 213. 

Type locality: Avera, Aroa River, Papua. 
Uromys siebersi Thomas, 1923, Treubia, 3: 422. 

Type locality: Gunung Daab, Great Kei Island. 

Nine specimens all from NE. New Guinea. Six from the Kratke Mts. : ^50.1233, 
$ 1234, Buntibasa district; <$ 1236, Apimuri; $ 1235, skull only 1238, Kambaidam; 
$ 1237, Sasara ; and three <$ 1678, 1679, x ^° from Degabaga, 8 miles east of Hagen 
Range, Sepik-Wahgi Divide. 

Macr uromys major Rummler 

Macruromys major Rummler, 1935, Z. Sdugetierk. 10: 105. 

Type locality: Buntibasa district, Kratke Mts., NE. New Guinea, 4,000-5,000 ft. 

Three specimens, all from NE. New Guinea: $ 50.1249, Saiko, Bubu River; 
<J 1250, Yampara, Kratke Mts. ; paratype $ 125 1 skull only, Buntibasa district, 
Kratke Mts. 

Lorentzimys alticola Tate & Archbold 

Lorentzimys nouhuysii alticola Tate & Archbold, 1941, Amer. Mus. Novit., No. 1101 : 4. 
Type locality: Nr. Lake Habema, Mt. Wilhelmina, Dutch New Guinea, 2,700 m. 

Eleven specimens. Six from NE. New Guinea: $ 47.1295, Baiyanka, SE. Bismarck 
Range ; $ 50.1730, $ 1732, juv. $ 1731, Yandara, high slopes Mt. Wilhelm ; <J 50.1728, 
$1729, high northern slopes Mt. Wilhelm, Bismarck Range; and five in alcohol 
from eastern Papua: $47.1296, 1298, juv. $1297, $1299, ? 1300, Enaena, Mount 
Simpson. 

The specimens from Baiyanka and Enaena were the first representatives of this 
genus to be received in London (Ellerman, 1949). The additional five specimens are 
very similar to these. 

Parahydromys asper (Thomas) 

Limnomys asper Thomas, 1906, Ann. Mag. Nat. Hist. 17: 326. 

Type locality: Mt. Gayata, Richardson Range, British New Guinea. 
Parahydromys Poche, 1906 (June), Zool. Am. 30: 326 (to replace Limnomys Thomas). 
Drosomys Thomas, 1906 (December), Proc. Biol. Soc. Washington, 19: 199 (to replace Limnomys 

Thomas) . 

Fifteen specimens, all from NE. New Guinea: (J 50.1240, 1241, 1242, 1243, Bunti- 
basa district, Kratke Mts. ; $ 1244, I2 45> Arau district, Kratke Mts. ; juv. # 1246, 

ZOO. I, IO T t 



3io 



MAMMALS COLLECTED BY MR. SHAW MAYER 



1247 (skull only), Kuraka, Kratke Mts. ; ^1248, Saiko, Bubu River; ^1669 (skin 
only), juv. $1670, Yanka, eastern slopes Hagen Range; juv. ^1671, Menebe, 8 
miles east Hagen Range, Sepik-Wahgi Divide ; $ 1672, Degabaga, 8 miles east 
Hagen Range, Sepik-Wahgi Divide; $1673, [juv. $1674 in spirit], Tomba, SW. 
slopes Hagen Range. 

Crossomys moncktoni Thomas 

Crossomys moncktoni Thomas, 1907, Ann. Mag. Nat. Hist. 20: 72. 
Type locality: Serigina, Brown River, NE. Papua, 4,500 ft. 

Fourteen specimens all from NE. New Guinea: ^50.1239, Arau, Kratke Mts.; 
S 1768, 1767, $ 1772, 1773, 1769, 1774, 1775, 1771, 1 77°> Baiyer River, east slope 
Hagen Range ; $ 1776, Tomba, SW. slopes Hagen Range ; <J 1777, $ 1778, 1779, 
Yandara, Bismarck Range. These are additional to the five specimens from Bai- 
yanka mentioned by Ellerman (1949). 

Leptomys elegans ernstmayri Rummler 

Leptomys ernstmayri Rummler, 1932, Das Aquarium 6: 131, 135. 
Type locality: Ogeramnang, Saruwaged Mts., Huon Peninsula, NE. New Guinea. 

Five specimens all from NE. New Guinea, ^50.1252, 1254 (skull only), $ 1253, 
1255 (skull only), Kambaidam, Kratke Mts. ; $ 1256, Arau district, Kratke Mts. 

Pseudohydromys murinus Rummler 

Pseudohydromys murinus Rummler, 1934, Z. Sdugetierk. 9: 48. 
Type locality: Morobe, Mt. Misim, NE. New Guinea, 7,000 ft. 

Three specimens, all from NE. New Guinea: (£50.1733, collector's No. 1136, 
Yandara, high slopes Mt. Wilhelm; $1734, collector's No. 1146, 1735, collector's 
No. 1 15 1, high northern slopes Mt. Wilhelm, Bismarck Range. 

These three specimens are new to our collection and appear to be the first speci- 
mens recorded since the type was described. The pelage agrees with the description 
of that of the type except that in No. 1734 it is a little shorter and greyer and in this 
same specimen the tip of the otherwise brown tail is white. The measurements also 
agree fairly well with those of the type and with the remeasurements by Tate (195 1) 
which show that the length of the nasals is 8-o mm. not 13.0 mm. as given by 
Rummler. 

These specimens extend the range of this species some 200 miles to the north-west 
of its type locality and to an altitude of 9,000-10,000 ft. 

Measurements in mm. (taken in the flesh) : 







* 








*Ǥ 




►«s 








►s; 








►0 












* 


<s 




« 

^ 


SP 








T3 
IS 






•X3 






1^ 


I* 


.3 


** — 

^ 




< 

>«<» 
« 


.3 




3 


CO 


fcq 




S 

s 




1 §p 
^ 


5o§ 


§ 


1 § 


3 




1 


1 


50.1733 


6* 


88 


91 


19 


9 


22-8 


io-6 


II'O 


47 


57 


2-1 


7'7 


3-1x1-1 


1734 


¥ 


103 


91 


20 


10 





io-8 


n-3 


4-5 


6-o 


2-1 


8-o 


2-8x0-9 


1735 


¥ 


105 


93 


19-5 


12 


22-9 


io-6 


ii-i 


4-8 


57 


2'0 


— 


3-2x1-1 



IN NEW GUINEA, 1932-1949 311 

Neohydromys gen. 

This is a small mouse-like Hydromyine, not modified for aquatic habits. It is 
distinguished from all other Hydromyinae, including Microhydromys which Tate & 
Archbold (1941) described as the smallest known Hydromyine, by having much 
smaller molar teeth, which are § as in most Hydromyinae, by its rather long muzzle 
which is short and broad in Microhydromys and by the large diastema which is 
larger than that of Microhydromys. The zygomatic plate is not so much excised in 
front as that of Microhydromys, and the upper incisor teeth are not grooved, a 
feature which appears to be unique to Microhydromys. The incisor teeth are, how- 
ever, well developed and are slightly pro-odont, as are those of Xeromys. The bullae 
are rather similar to those of the type of Microhydromys (measurements of the 
type of Microhydromys are given in parentheses): width 2-9 mm. (2-9 mm.), length 
4-0 mm. (3-8 mm.), distance apart 2-2 mm. (2.0 mm.). The palatal foramina are 
small as in Pseudohydromys, but the pterigoid and alisphenoid region is not 
swollen. The angular projection of the mandible is not so pronounced as in Pseudo- 
hydromys. 

Type species: Neohydromys fuscus 

Neohydromys fuscus sp. n. 

Type locality: High northern slopes Mt. Wilhelm, Bismarck Range, NE. New 

Guinea, 9,000-10,000 ft. 
Type: Adult $ 50.1736, collector's No. 1185, 19 June 1949. Skin and skull. 

In external appearance this small murid is very similar to Pseudohydromys murinus. 
(I have not seen a specimen of Microhydromys richardsoni with which it also appears 
to be very similar in external appearance.) The pelage, which is about 4 mm. long, 
is smoky grey in colour and only slightly lighter ventrally. The ears are the same 
colour as the body. The fore and hind feet are slender and lightly covered with short 
white hairs. The tail is brownish both above and below ; according to the collector 
the terminal 16 mm. was white ; there are 17 rings of scales per centimetre, and the 
fine silvery scale hairs are only about half the length of the scale. The skull is a little 
larger than that of Pseudohydromys murinus but is easily distinguished from it by 
the very small molars, the slightly pro-odont incisor teeth (upper ones pale orange 
with white tips, lower ones pale yellow), the longer muzzle and larger diastema, and 
the less excised zygomatic plate. Neohydromys fuscus is also distinct from Pseudo- 
hydromys occidentalis Tate (195 1). 

Measurements in mm. (taken in the flesh) : 
Skin: head and body 92 ; tail 78 ; hind foot 21 ; ear 12. 

Skull: condylo-basal length 24-3; zygomatic breadth 12-3; palatal length 13-1; 
inter-orbital breadth 5-2; diastema 8-2; palatal foramina 2-0; nasals (length) 
7-9; bullae, 4-0x2-9; distance apart of bullae, 2-0; palatal breadth between 
n^-m 1 2-6; length m 1 +m 2 2-1; m 1 (length x breadth) 1-4x0-7; m 1 (length x 
breadth) 0-7x0-6; mandible, greatest length (except incisors), 13.4; m 1 + m 2) 
2-2. 
zoo. 1, 10 t t 2 



312 



MAMMALS COLLECTED BY MR. SHAW MAYER 
CHIROPTERA 



Pteropus sp. 

One young (J 50.969 (skin only) from Fergusson Island. 

Dobsonia moluccensis magna Thomas 

Dobsonia magna Thomas, 1905, Ann. Mag. Nat. Hist. 16: 423. 

Type locality: Tamata, Mambare River, eastern New Guinea. 
Dobsonia moluccensis magna Thomas, Andersen, 1912, Cat. Chiropt, Coll. Brit. Mus., 2nd ed., 

1. Megachiroptera, 825, London. 

Two specimens, <J 50.1149, 1150, Buntibasa district, Kratke Mts., NE. New Guinea. 

Nyctimene papuanus Andersen 

Nyctimene papuanus Andersen, 1910, Ann. Mag. Nat. Hist. 6: 621. 
Type locality: Milne Bay, eastern tip of New Guinea. 

One specimen c? 50.1153, Arau district, Kratke Mts., NE. New Guinea. 
Paranyctimene raptor Tate 

Paranyctimene raptor Tate, 1942, Amer. Mus. Novit., No. 1204: 1. 
Type locality: Oroville Camp, Fly River, Papua. 

Two specimens collected 12 January 1933 which are new to our collection: 
550.1151, $ 1152 from the Arau district, Kratke Mts., NE. New Guinea. 

I have not been able to compare these specimens with the type but they appear 
to be very similar to the description of it ; they have no dorsal stripe. Some of their 
measurements are a little larger but those for the teeth agree closely with those of the 
type. 

Measurements in mm. (taken in the flesh) : 



























<» 








?s 




















<s 








^ 














to 






<^> 


^ 















<y> 












s 


^ 






^s 




^» 
















^ 


Q 


8 
<» 




00 


1 


s 


5 


** 
« 

« 


to 

1 


1 



^ bo 


~ 


►0 


5 


3 
5 


50-1151 


6* 


77 


19 


— 


10 


4 


55 


25*2 


4'5 


16-2 


4-2 


10-7 


12-9 


1152 


¥ 


80 


20 


— 


10 


4 


55 


— 


4'5 


16-2 


5-o 


II-O 


— 

















<» 
8 


8 








-£ 






1 


"So 




"So 




i 


•<s» 












*>. <o 


is 
li 


§ 
^ 





.3 







^ § 





§ 
^ 







^ 


■§.§ 




*««0 




<^<o 


5 $ 


•^CO 


§ 

S 




S 


.§ 


•s ^ 


^1 


s 


^a 


s 


3.2 






sa 


S 


§ 


£ 

$ 




tuow 

8 -Si 


■SP§ 
as « 




•Sp§ 


S 


•^§ 


^^ 


S 2 


? 


* 


k « 


^ 


0^ 


^1 




fea 


tq « 


O^ 




O 


50.1151 


4.8 


i'3 


1-7 


2-2 


i-6 


1-8 


3'7 


1-2 


2-6 


17 


2-0 


1-7 


1152 


4'7 


i*3 


1-7 


2-2 


i-6 


i'5 


3-8 


i*3 


2*5 


1-7 


2-0 


1-7 



IN NEW GUINEA, 1932-1949 313 

Syconycteris crassa papuana (Matschie) 

Macroglossus {Syconycteris) papuanus Matschie, 1899, Die Fledermdnse des Berliner Museums 
f. Naturkunde, Megachiroptera: 99. Berlin. 
Type locality: Andai, NW. New Guinea. 
Syconycteris crassa papuana (Matschie), Andersen, 1912, Cat. Chiropt. Col. Brit. Mus., 2nded., 
1. Megachiroptera, 777, London. 

Six specimens. Five from NE. New Guinea: $ 50.1798, 1799, Yandara, Bismarck 
Range, $970 [in spirit], Baiyanka, SE. Bismarck Range; $1800, Juzaing, Sara- 
waged Range, Huon Peninsula; $ 1801 [in spirit], Tomba, Hagen Range, and one 
juv. $ 971 [in spirit], from Enaena, NE. slopes Mt. Simpson, eastern Papua. 

The spirit specimen from Tomba differs from the others by its narrow teeth, and 
it has less hair on the forearm than in the average specimens. 

Hipposideros muscinus muscinus (Thomas & Doria) 

Phyllorhina muscina Thomas & Doria, 1886, Ann. Mus. Stor. nat. Genova, 24: 203. 

Type locality: Fly River, Papua. 
Hipposideros muscinus muscinus (Thomas & Doria), Tate, 1946, Amer. Mus. Novit., No. 
1323: 1-21. 

Two specimens, ^50.1154, $1155, Buntibasa district, Kratke Mts., NE. New 
Guinea. 

Philetor rohui Thomas 

Philetor rohui Thomas, 1902, Ann. Mag. Nat. Hist. 9: 220. 
Type locality: Albert Edward Range, Papua, 6,000 ft. 

Seven specimens in spirit : <$ 50.972, 973, $ 974, 975 976, 977, 978 from Enaena, 
NE. slopes Mt. Simpson, eastern Papua. 

Pipistrellus collinus Thomas 

Pipistrellus papuanus collinus Thomas, 1920, Ann. Mag. Nat. Hist. 9: 533. 

Type locality : Bihagi, head of Mambari River, Papua. 
Pipistrellus collinus Thomas, Tate, 1942, Bull. Amer. Mus. Nat. Hist. 80: 241. 

One $ 50.983 [in spirit], Baiyanka, SE. Bismarck Range, NE. New Guinea. 

Miniopterus schreibersi blepotis (Temminck) 

Vespertilio blepotis Temminck, 1841, Monographies de Mammalogie. ... 2: 212. Paris 6- 
Amsterdam. 
Type locality: Java — also Banda, Amboina, Timor, Japan. 
Miniopterus schreibersii blepotis Temminck = medius = ravus — eschscholtzii = fuscus = 
yayeyamae) Tate, 1941, Bull. Amer. Mus. Nat. Hist. 78: 567-597. 

Two specimens, $50.1802, 1803 [in spirit], Tomba, Hagen Range, NE. New 
Guinea. 



314 MAMMALS COLLECTED BY MR. SHAW MAYER 

Miniopterus schreibersi magnater Sanborn 

Miniopterus schreibersi magnater Sanborn, 1913, Field Mus. Publ. Zool. 18: 26. 
Type locality : Sepik River, New Guinea. 

Three specimens, $ 50.1156, 1158, $ 1157, Arau district, Kratke Mts., NE. New 
Guinea. 



Otomops secundus sp.n. # 

Type locality: Tapu, Upper Ramu River Plateau, NE. New Guinea. 
Type: Adult S 50.982, collector's No. 568 [in spirit]. 

Paratypes: $ 50. 979, collector's No. 565, 980, collector's No. 566, 981, collector's 
No. 567. All in spirit (skulls extracted). 

Since the recent (1948) discovery of the remarkable genus Otomops in New Guinea 
(0. papuensis Lawrence, type locality Vailala River, western Papua) these are the 
first additional specimens to be collected. While they are no doubt closely related 
to 0. papuensis, their considerably longer forearm and well-marked pale mantle 
make it necessary to recognize them as distinct. 

It is a small Otomops with all the distinctive external and cranial characters of 
the genus ; with forearm 57 (type) and 58 mm. in length (49*2 in the type and only 
specimen of 0. papuensis) and with broad pale buffy-grey mantle as in 0. wroughtoni 
and other species. Colour: dark chocolate-brown on nape and lower back, darkest 
on lower back. Crown pale brown. Mantle across shoulders well defined, especially 
anteriorly, and consisting of pale buffy or greyish hairs of which only a few have 
dark tips. Along the margin of the membranes adjoining the body, above, there is 
a conspicuous but narrow white line composed of very short pure white hairs sharply 
outlining the deep chocolate of the body colour. 

As in 0. papuensis, the premaxillaries are open. Little importance should be 
attached to this feature, however, since in a series of eleven skulls of 0. wroughtoni, 
type species of the genus, two have the premaxillaries separated, although their 
union was said by Thomas to be one of the generic characters. In the very deep 
basisphenoid pits and in the forward extension to the pterygoids of the tympanic 
bullae, as well as in the extension of the zygomatic plate, this new form presents 
(as Lawrence remarks of 0. papuensis) characters of greater generic value than open 
or closed premaxillae. 

Otomops secundus differs from 0. formosus Chasen of Java in much smaller skull, 
2i-2 (against 24), although the forearm measurements of the two forms are closely 
approximate (59-7 in 0. formosus). Although the forearm in 0. secundus is nearly 
10 mm. longer than in 0. papuensis, the cranial measurements show little difference. 
The type locality of the new form is little more than 100 miles north of that of 0. 
papuensis, but is separated from it by the central mountain range. It is possible 
that further collecting in New Guinea and other parts of the Indo-Australasian 
Archipelago may eventually bring to light intermediate forms and so reduce to 

* The description of this species is by Mr. R. W. Hayman. 



IN NEW GUINEA, 1932-1949 



315 



subspecific rank some of the named species; but until then it seems advisable to 
separate specifically the present form. 

Measurements in mm. (External from spirit specimens: type of 0. papuensis in 
parentheses) : 







* 






Si 














-o 






« 






< 


Q 


s 






I*"* 



8 



1-3 


^. 


S 






1 




^ 


« 

h 


s 


sa 


« 

^ 


O 


si 


s 


50.982 Type 


a 


71 (67) 


37 (3o) 


10 (io-6) 


23-9 (23-5) 


24 (22-2) 


58 (49'2) 


21-2 (20-2) 


19*3 


979 


¥ 


68 


38 


10 


22-0 


24'3 


58 


21-0 


19-9 


980 


¥ 


70 


36 


10 


24-9 


24 


57 


21-5 


19-8 


981 


$ 


68 


33 


10 


24-6 


23 


58 


2I-0 


19-5 





























^ 




to 














s 


•^ 




O 










1 

1 


t>jo « 

N -0 


.■0 

s 


■a! 







f 




CO 

I 


n 

? 

<«5 


50.982 Type 


II-2 


10-9 (9-5) 


9-9 (9'9) 


i3-i 


4-5 


8-o 


8-6(8-5) 


7'5 


8-o (7.7) 


979 


II-I 


II-2 


9-9 


14-0 


4-2 


8-o 


8-7 


7-6 


8-i 


980 


II-O 


II-O 


9-9 


14-0 


4.4 


8-o 


8-7 


7-8 


8-2 


981 





10-9 


IO-I 


13-7 


4-4 


8-o 


8-7 


7-8 


8-o 



REFERENCES 

Carter, T. D., Hill, J. E., & Tate, G. H. H. 1945. Mammals of the Pacific World, pp. xvi 
+ 227, text figs. New York. 

Dollman, G. 1930. On mammals obtained by Mr. Shaw Mayer in New Guinea, and pre- 
sented to the British Museum by Mr. J. Spedan Lewis, F.Z.S. Proc. Zool. Soc. Lond. 1930: 

429-435. 

Ellerman, J. R. 1940, 1941, and 1949. The Families and Geneva of Living Rodents. Vols. 

1, 2, and 3 respectively. London. 
Hinton, M. A. C. 1943. Preliminary diagnosis of five new murine rodents from New Guinea. 

Ann. Mag. Nat. Hist. 10: 552-557. 
Jentink, F. A. 1908. Mammals collected by the members of the Humboldt-Bay and the 

Merauke River-expeditions. Nova Guinea. Uitkomsten der Nederlandsche Nieuw-Guinea 

Expeditie in igoj onder hiding van. . . . A. Wichmann, 5: 361-364, Leiden. 
191 1. New and interesting mammals of the Dutch New-Guinea-Expedition to the Snow 

Mountains. Notes Ley den Mus. 33: 233-238. 
Le Souef, A. S., & Burrell, H. 1926. The Wild Animals of Australasia. . . . with a chapter on 

Bats by Ellis le G. Troughton. pp. 387, 57 pis., London. 
Miklouho-Maclay, N. de. 1884. Notes on zoology of the Maclay-coast (1) in New Guinea. 

Proc. Linn. Soc. New South Wales, 9: 713-720. 
Rummler, H. 1935. Neue Muriden aus Neuguinea. Z. Sdugertierk. 10: 105-118. 
1938. Die Systematic und Verbreitung der Muriden Neuguineas. Mitt. Zool. Mus. Berlin, 

23: 1-297- 



3 i6 MAMMALS COLLECTED BY MR. SHAW MAYER 

Schwarz, E. 1934. On a wallaby and a phalanger brought by Mr. Wilfred Frost from the 
islands west of New Guinea. With notes on the evolution of coat, colour, and pattern in 
the genus Phalanger. Proc. Zool. Soc. Lond. 1934: 87-91. 

Tate, G. H. H., & Archbold, R. 1937. Results of the Archbold Expeditions, 16. Some 
marsupials of New Guinea and Celebes. Bull. Amer. Mus. Nat. Hist. 73: 331-476. 

■ 1 94 1. Results of the Archbold Expeditions, 31. New rodents and marsupials from 

New Guinea. Amer. Mus. Novit., No. 1101: 2. 

1 94 1. Results of the Archbold Expeditions, 35. A review of the genus Hipposideros with 

special reference to Indo-Australian species. Bull. Amer. Mus. Nat. Hist. 78: 353-393. 

1 94 1. Results of the Archbold Expeditions, 38. Molossid bats of the Archbold collec- 
tions. Amer. Mus. Novit., No. 1142: 1-4. 

1941. Results of the Archbold Expeditions, 40. Notes on vespertilionid bats of the sub- 
families Miniopterinae, Murininae, Kerivoulinae, and Nyctophilinae. Bull. Amer. Mus. Nat. 
Hist. 78:567-597. 

1942. Results of the Archbold Expeditions, 47. Review of the vespertilionid bats, with 

special attention to genera and species of the Archbold collections. Bull. Amer. Mus. Nat. 
Hist. 80: 221-297. 

1942. Results of the Archbold Expeditions, 48. Pteropodidae (Chiroptera) of the Arch- 
bold collections. Bull. Amer. Mus. Hist. 80: 331-347- 

1945. Results of the Archbold Expeditions, 52. The marsupial genus Phalanger. Amer. 

Mus. Novit., No. 1283: 1-41. 

1945. Results of the Archbold Expeditions, 54. The marsupial genus Pseudocheirus and 

its subgenera. Amer. Mus. Novit., No. 1287: 1-30. 

1945. Results of the Archbold Expeditions, 55. Notes on the squirrel-like and mouse-like 

possums (Marsupialia). Amer. Mus. Novit., No. 1305: 1-12. 

1946. Geographical distribution of the bats in the Australasian Archipelago. Amer. Mus. 

Novit., No. 1323: 1-2 1. 

1947. Results of the Archbold Expeditions, 56. On the anatomy and classification of the 

Dasyuridae (Marsupialia). Bull. Amer. Mus. Nat. Hist. 88: 97-156. 

1948. Results of the Archbold Expeditions, 59. Studies on the anatomy and phylogeny 

of the Macropodidae (Marsupialia). Bull. Amer. Mus. Nat. Hist. 91: 233-352. 

1948. Results of the Archbold Expeditions, 60. Studies in the Peramelidae (Marsupialia). 

Bull. Amer. Mus. Nat. Hist. 92: 313-346. 

1951. Results of the Archbold Expeditions, 65. The rodents of Australia and New Guinea. 

Bull. Amer. Mus. Nat. Hist. 97: 183-430. 

Thomas, M. R. O. 1888. Catalogue of the Marsupialia and Monotremata in the ... British Museum, 
pp. xiii+401, 28 pis. (col.), London. 

■ 1920. New small mammals from New Guinea. Ann. Mag. Nat. Hist. 6: 533-537. 



IN NEW GUINEA, 1932-1949 317 

APPENDIX I 

LIST OF LOCALITIES 
FROM WHICH SPECIMENS WERE OBTAINED 

North-East New Guinea 

Apimuri (Buntibasa district), Kratke Mts., 4,500 ft. 

Arau district, Kratke Mts., 4,000-5,500 ft. 

Baiyanka, Purari-Ramu Divide, SE. Bismarck Range, 7,500-8,500 ft. 

Baiyer River, nr. Yanka, east slopes Hagen Range, Central Highlands, 8,000 ft. 

Binemarian, Kratke Mts., 4,000-5,000 ft. 

Bogo, 50 miles east of Hagen Government Station, south slopes Bismarck Range, 6,000 ft. 

Bubu River district (Upper Waria River), 5,000-8,000 ft. 

Buntibasa district, Kratke Mts., 4,000-5,500 ft. 

Degabaga, 8 miles east of Hagen Range, 25 miles north of Hagen Govt. Station, Sepik-Wahgi 

Divide, Central Highlands, 4,500-6,000 ft. 
Garaina, Upper Waria River, 2,500-3,000 ft. 

Guyebi, northern slopes Mt. Wilhelm, Bismarck Range, 6,000-7,000 ft. 
Herowagi (42 miles east of Hagen Govt. Station), south slopes Bismarck Range, 7,000 ft. 
High slopes Mt. Wilhelm, Bismarck Range, 9,000-10,000 ft. 
Junzaing, Saruwaged Range, Huon Peninsula, 6,000 ft. 
Kambaidam (Buntibasa district), Kratke Mts., 4,000 ft. 
Kuraka (Buntibasa district), Kratke Mts., 4,000-5,000 ft. 
Menebe, 8 miles east of Hagen Range, 20 miles north of Hagen Govt. Station, Sepik-Wahgi 

Divide, Central Highlands, 4,500-5,500 ft. 
Mendi, northern slopes Mt. Wilhelm, Bismarck Range, 4,500 ft. 
Saiko, Bubu River (Upper Waria River), 5,000-7,000 ft. 
Sasara (Buntibasa district), Kratke Mts., 4,500-5,500 ft 
South and north side Bubu River (Upper Waria River), 6,000-7,000 ft. 
Tapu, Upper Ramu River Plateau, 6,000 ft. 

Tomba, south-west slopes Hagen Range, Central Highlands, 8,000-9,500 ft. 
Yampara (Buntibasa district), Kratke Mts., 4,700 ft. 

Yandara, northern slopes Mt. Wilhelm, Bismarck Range, 5,500-10,000 ft. 
Yanka, eastern slopes Hagen Range, Central Highlands, 5,000-8,000 ft. 
Zageheme, Cromwell Mts., Huon Peninsula. 

Eastern Papua, South-East New Guinea 
Bibitau, Mt. Orian (30 miles NW. Mt. Simpson), Main Range, 2,500 ft. 
(Boneno Camp), Mt. Maneao (35 miles NW. Mt. Simpson), Main Range, 6,000 ft. 
Boneno, Mt. Mura (30 miles NW. Mt. Simpson), c. 4,000-7,000 ft. 
Enaena, NE. slopes Mt. Simpson, 1,000-6,500 ft. 
Ikara, NE. slopes Mt. Simpson, 3,500-5,000 ft. 
Maneao Range (35 miles NE. Mt. Simpson), 7,000 ft. 
Mt. Mura (30 miles NW. Mt. Simpson), Main Range, 5,000 ft. 
Wapona, north slopes Maneao Range (35 miles NW. Mt. Simpson), 1,000 ft. 

Other Localities 

Faralulu district, West Fergusson Island, SE. New Guinea, 600 ft. 

Taibutu district, West Fergusson Island, SE. New Guinea, 900-1,100 ft. 

Mountains above Taibutu village, West Fergusson Island, SE. New Guinea, 2,000-3,000 ft. 

Lau, Bainings Mts., Gazelle Peninsula, New Britain, 1,300 ft. (Rattus exulans browni only.) 

Mountains SE. New Guinea, behind island of Samaria. 

Tongoa Island, New Hebrides, 400 ft. (Rattus exulans exulans only.) 



3 i8 MAMMALS COLLECTED BY MR. SHAW MAYER 

APPENDIX II 

FORMS DESCRIBED AS NEW IN THIS PAPER 

Zaglossus bubuensis 

Dactylopsila tatei 

Pseudocheirus (Pseudochirops) corinnae fuscus 

Peroryctes longicauda magna 

Peroryctes papuensis 
Murexia longicaudata parva 

Antechinus hageni 
Pogonomys fergussoniensis 

Pogonomys shawmayeri 

Rattus ruber fergussoniensis 

Rattus verecundus tomba 

Neohydromys fuscus (new genus) 

Otomops secundus 



APPENDIX III 
Dendrolagus dorianus notatus Matschie 

Dendrolagus notatus Matschie, 1916, Mitt. zool. Mus. Berlin, 8: 294. 

Type locality : Slopes of the Schrader Mountains, between 5 S. and 144 E., NE. New Guinea. 

Two specimens, a young adult ^50.1815 and a juv. ? 1816 from Yanka, eastern 
slopes Hagen Range, 8,000 ft. 

These specimens were collected about 30 miles away from the type locality and 
appear to be the first to be recorded since the type was described from a single 
specimen. 

Measurements in mm. (taken in the flesh) : 















* 


















.0 
§ 




s 

&j5 


§ 




s 
&3 






Is 


1 


§ 


3\ 


s 


1 


"g 


§ 


50.1815 


* 


610 


470 


108 


50 


l65'5 


66-2 


4i-8x 21-0 


13-8 


io-6x 6-o 


7-0 x 6-i 


7-0 x 6-6 


6-8x6-6 


6-5 X 6-3 




1 DEC 1952 



PRINTED IN 

GREAT BRITAIN 

AT THE 

UNIVERSITY PRESS 

OXFORD 

BY 

CHARLES BATEY 

PRINTER 

TO THE 

UNIVERSITY 




2 8 JAN 1953 



TAXOTIOMY OF THE 

KARROO AND 

RED-BACK LARKS 

OF WESTERN 

SOUTH AFRICA 

J. D. MACDONALD 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 

ZOOLOGY Vol. i No. ii 

LONDON: 1953 



• *te\ 






TAXONOMY OF THE KARROO AND 

RED-BACK LARKS OF 

WESTERN SOUTH AFRICA 



BY 

J. D. MA CDONAL 1 



Mj. 




Pp. 319-350; Pis. 36-38; 5 Text-figures 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ZOOLOGY Vol. 1 No. 11 

LONDON: 1953 



THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted in ig4g , is issued 
in five series, corresponding to the Departments of the 
Museum. 

Parts appear at irregular intervals as they become 
ready. Volumes will contain about three or four hundred 
pages, and will not necessarily be completed within one 
calendar year. 

This paper is Vol. i, No. II of the Zoological series. 



PRINTED BY ORDER OF THE TRUSTEES OF 
THE BRITISH MUSEUM 

Issued January ig$3 Price Ten Shillings 



TAXONOMY OF THE KARROO AND RED- 
BACK LARKS OF WESTERN SOUTH AFRICA 

By J. D. MACDONALD 

[Received ist September 1951] 



321 
321 
323 
323 



Introduction ......... 

Historical Note ......... 

Methods . . . . . . 

Materials and Acknowledgements . . . . . 

Populations Examined: 

Cape Flats . . . . . . . . ' . . 324 

Berg River and Saldanha Bay ....... 325 

Lambert's Bay .......... 325 

Swellendam and Deelfontein ....... 326 

Traka and Nels Poort ......... 326 

Namaqua-Bushman-land Region ....... 327 

Orange River Mouth ......... 329 

Witputs Area .......... 330 

Aus Area ........... 331 

Namib Dunes .......... 333 

Discussion 

Dimensions: Colour and Pattern: Developmental Stages: Breeding- 
cycle: Habits: Nomenclature ....... 335 

Summary ........... 345 

References ........... 345 

Tables of Measurements ........ 346 

SYNOPSIS 

A review of the Karroo and Red-back Larks in the light of recent data indicates that these two birds, 
long regarded as separate species, and sometimes placed in different genera, are in fact geographical races 
of the same species. Variation is limited almost entirely to colour and pattern, and there is a gradual transi- 
tion from one extreme form to another. Seven geographical races are recognized, two of which are new. 

INTRODUCTION 
Material and data collected by the British Museum (Natural History) South West 
Africa Expedition (1949-1950) throw new light on the taxonomy of the Karroo and 
Red-back Larks, usually assigned to the genera Calendulauda and Pseudammo- 
manes. Birds of two different colours, apparently identical in every other respect, 
were found in each other's company near the mouth of the Orange River. The 'grey' 
birds were thought to be Calendulauda albescens and the 'red' birds Pseudammo- 
manes ( ? species). It now seems that these birds belong to two groups which may be 
more closely related than has been recognized hitherto. The purpose of this paper is 
to examine this matter in the light of the data now available. 

HISTORICAL NOTE 
The published history of these larks begins with the description of the Karroo 
Lark by Lafresnaye 1 (1839:259). He described two species, first Alauda albescens 

1 The name Certhilauda nivosa Swainson, 1837, though often used for the Karroo Lark, was shown 
by Roberts (1936a: 257) to be inapplicable, being based on a juvenile Galerida cristata senegalensis. 



322 TAXONOMY OF THE KARROO AND 

from Blauw-Berg (Blaauwberg Beach, in the north of Table Bay), a 'grey' bird, 
and then a 'red* species A. guttata from Elephant's River (now Oliphant's River), 
Cape Province. The types are in the Museum of Comparative Zoology, Cam- 
bridge, Mass., U.S.A.: that of A. guttata is said to be a bird in juvenile plumage. 
Unaware of Lafresnaye's description Andrew Smith (1843) also described 'grey* 
and 'red' birds as separate species, the former as Alauda codea and the latter as A. 
lagepa, both of which were figured together on Plate 87. Specimens of each of 
his species are represented in the National Collection, but they can only be 
regarded as co-types, for Smith did not designate types and his collection was 
split up. 

An analysis of Smith's descriptions shows that in dimensions, distribution, and 
habits these two species are very similar. Sharpe (1874: 624) came to the conclusion 
'that they are nothing but the summer and winter plumage of the same bird. However 
curious this may seem, I think it is not to be refuted on the evidence of the speci- 
mens which I have before me'. 

Sharpe's opinion held until Roberts (19360: 258; 1940: 191) revived the concep- 
tion of two sympatric species differing mainly in colour and divisible into several 
geographical races. In the pallid or grey form, Calendulauda albescens, he recog- 
nized three races, the typical one, a second race C. a. saldanhae, with a strong 
tinge of rufous on the upper parts, but not so rufous as C. guttata, and a third C. a. 
karruensis, very dark above with some rufous in parts. In the rufous form, C. 
guttata, he recognized two races, C. g. calviniensis being slightly larger than the 
typical form. 

Meinertzhagen 1 (195 1: 107) put all these variations into one polymorphic species, 
C. albescens, in which variation could not be correlated with distribution. He calls 
it 'a very variable bird in both size and colour throughout its range. It has a pure 
grey and a pure red phase, with every intermediate and without constancy in 
distribution'. 

The first Red-back Larks were found by Andersson in the dry sandy bed of the 
Kuiseb River, near Walvis Bay. Strickland (1852) named them Alauda erythroch- 
lamys. Several birds found in other localities, notably in the Transvaal by Ayres 
(1874), were wrongly associated with this species, but remained with it until removed 
by Roberts. In fact the true Red-back Lark does not seem to have been recorded 2 
again until Roberts (1937 : 95) found it about 30 miles north of Aus, a place about 
60 miles inland from Luderitz Bay and about 300 miles south of Walvis Bay. Roberts 
also found specimens nearer Aus, but concluded that they belonged to a different 
species, which he named Pseudammomanes barlowi. 

Hoesch and Niethammer (1940: 224) did not agree with Roberts and maintained 
that the Aus birds were inseparable from those at Walvis Bay. Neither did Meinertz- 
hagen (1951: 107), but he put the species into the genus Certhilauda along with the 
Karroo Lark and several other species. 

1 I am indebted to Colonel R. Meinertzhagen for his courtesy in lending me a typescript copy of his 
paper which was in course of publication when this paper was in preparation. 

2 R. D. Bradfield collected a specimen on the Kuiseb River on the 18th of December 1928, but it 
does not seem to have been recorded. It is in the Transvaal Museum. 



RED-BACK LARKS OF WESTERN SOUTH AFRICA 323 

METHODS 

The morphological characters examined here are lengths of wing, tail, bill, and 
first primary; also colour and colour-pattern. Other characters considered are 
moults, breeding-cycle, developmental stages, and habits. 

Wing measurements are taken on the stretched wing ; tail measurements from the 
crotch of the two central tail feathers, into which the leg of a divider can be firmly 
pressed, to the tip of the longest feather ; bills are measured from the cranio-f acial 
angle to the tip ; and first primary from the hard sheath covering the base of the 
shaft. The degree of error due to the set of the wing, the variation in the method 
of stretching, and the age of the critical feathers, whether fresh or not quite fully 
grown, or in various stages of wear, made it impossible to attempt extremely accurate 
measurements. The purpose of the measurements is merely to discover general 
correlations and they are taken to the nearest millimetre. 

For standards of colour Villalobos's Colour Atlas (1947) was used. This atlas 
contains a range of thirty-eight hues each of which is divided into a number of tones 
obtained by the combination of two variables, degree of lightness and degree of 
chromaticity. For example, in the symbol OOS/12/5 the OOS indicates the hue 
which is a mixture of two parts orange and one part scarlet ; the figure 12 indicates 
the degree of lightness, the range 1-20 being from darkest to lightest ; and the figure 
5 the degree of chromaticity, the range 1-12 being from the least to the greatest 
intensity of colour. Even without the Colour Atlas these symbols can convey some 
meaning to the reader, at least in a comparative sense. For example, of the differently 
coloured larks of this group under examination from near the mouth of the Orange 
River the 'grey' birds match approximately OOS/9/4 and the 'red' birds OOS/8/5 ; 
the inference is that they belong to the same colour group, or hue, but that the 
'grey' birds are one degree lighter and one degree less colourful than the 'red' birds. 

The use of these symbols may be regarded as an experiment in this method of 
colour determintation and colour comparison. The conclusions reached so far is 
that, though it is not ideal, it is unquestionably more satisfactory than the usual 
descriptive terminology, which often means one thing to the writer and something 
quite different to the reader. The Colour Atlas gives a cross-reference to colour names 
in common use, and where possible these have been included in the text. In birds 
with a streaked pattern the coloured area referred to in the following notes is that 
found outside the dark centres of the feathers (see Fig. 4). 

MATERIALS AND ACKNOWLEDGEMENTS 

This study is based on 80 specimens collected by the British Museum (Natural 
History) South West Africa Expedition (1949-1950), 36 other specimens in the 
National Collection, 45 in the Transvaal Museum, Pretoria, 13 in the South African 
Museum, Cape Town, and 21 in the private collection of Colonel R. Meinertzhagen. 

For their kindness in giving me permission to examine specimens, and sending 
others to me for examination, I have to thank Dr. V. FitzSimons and Mrs. J. Camp- 
bell, of the Transvaal Museum; Dr. K. Barnard, of the South African Museum; and 
Colonel R. Meinertzhagen. 



324 TAXONOMY OF THE KARROO AND 

I am indebted to Mr. J. D. M. Keet, of the Department of Agriculture, Pretoria, 
for information on the Aristida grasses of the Namib. For obtaining permission to 
enter the diamond controlled area at Tsondab Mund to look for these larks I am 
indebted to Mr. A. D. Vos, Inspector of Mines, Windhoek; and to Colonel Mentz, 
of the South African Police, for providing us with a police escort. 

Many of the specimens and data obtained by the Expedition were collected by 
two of my companions, Colonel F. O. Cave and Mrs. B. P. Hall. As a small tribute 
to their assistance, two new geographical races, based on material collected by the 
expedition, have been named after them. 

POPULATIONS EXAMINED 

In the first instance the evidence of specimens in various localities and areas will 
be examined: these places are located on the map, Plate 36. It is convenient to begin 
with the Cape area. 

Cape Flats 

Eleven specimens from localities in the Cape Flats have been examined. They 
are from Blaauwberg, on the coast about 10 miles north of Cape Town, the type 
locality of Alauda albescens) Milverton, a few miles north of Blaauwberg; Durban- 
ville, about 10 miles out of Cape Town on the Wellington road ; and Philadelphia, 
about 20 miles out on the Malmesbury road. There is also an old Butler specimen 
labelled 'Cape Town'. All these birds are similarly 'grey' in colour, actually a light 
drab, about OOS/11/3 in the Colour Atlas. Birds of this colour have not been re- 
corded from localities outside the Cape Flats, other than 'grey' birds of a slightly 
different tone which occur along the coast (see notes on Berg River and Saldanha 
Bay specimens). But three Smith specimens which are identical and belong to his 
'grey' A . codea require a special note, for Smith (1843) gives the range of this species 
as 'generally found upon the Karroo plains between the Oliphant and Orange 
Rivers'. Roberts (19360 : 312) shows that Smith had made entries in his early diaries 
on a lark found in the vicinity of Cape Town which, as Roberts points out, might 
easily have been this species. When he prepared his Zoology some years later, Smith 
must have had some difficulty in sorting out his data referable to birds he then 
described as the new species, A. codea and A. guttata, especially as these birds now 
appear to be polychromatic variations of the same species. It is my opinion, therefore, 
that Smith actually obtained the specimens on which he based his A. codea in the 
Cape area and not between the Oliphant 's and Orange rivers. 

One of Smith's specimens is just completing moult. It is undated, but may be 
the specimen referred to by Smith in his notes, see Roberts (19366: 313), 'on the 
4 December killed a young lark ... on the ascent of the Lions Rump'. 

According to A. W. Vincent (1946: 446) 'they begin to show breeding activity in 
early August' ; he has seen young birds about in October, but also nests with eggs 
in November. He says that this lark 'appears to be confined to the lower shady 
ground to the northward along the coast and close to the shores, becoming common 
farther out and extending through the drier western districts'. 



RED-BACK LARKS OF WESTERN SOUTH AFRICA 

The dimensions of the fourteen specimens referred to are as follows : 



325 



Sex 


No. 


Wing 


Tail 


Bill 


F.P. 


? 
? 


5 
4 
5 


88-94 

82-88 
85-90 


62-68 
56-61 
60-64 


18-20 
18-20 
17-19 


28-32 
28-33 
27-33 



The Cape Flats population, therefore, seem to have a distinctive light drab colour 
and the small range of measurement shows that females are slightly smaller than 
males. The main breeding-period appears to be about September to November, and 
the usual habitat is sandy scrub. 

Berg River and Saldanha Bay 

One Layard specimen from Berg River matches the Cape specimens perfectly 
in colour. Two other Layard specimens from the same locality have a slight pinkish- 
rufous wash on the upper parts, about OOS/10/3. All are undated and only one is 
sexed. Modern maps show Berg River as a locality about 70 miles north of Cape 
Town and about 15 miles from the mouth of the Great Berg River which opens into 
St. Helena Bay. On old maps the river itself is so named and there is therefore no 
certainty that the specimens were collected in exactly the same locality. 

Roberts (1936a: 258) found rufous-coloured birds at Saldanha Bay, which is on 
the coast about 20 miles west of Berg River and 60 miles north of Cape Town. He 
described them as having 'a strong wash of rufous on all the upper parts, but not as 
rufous as in Calendulauda guttata' . Two specimens collected by Shortridge in 1903 
at Hoetzes Bay, Saldanha Bay, which are in the South African Museum, fit this 
description and are exactly similar to our pinkish-rufous Berg River specimens. 
They are rather worn. One of Roberts's specimens, collected in November, is in 
juvenile plumage. 

Measurements of Berg River and Saldanha Bay specimens are as follows: 



Sex 


No. 


Wing 


Tail 


Bill 


F.P. 


? 

? 


3 
1 

2 


9i 

88 

90-93 


62-68 

61 
64-66 


19-20 
20 
20 


31-32 

27 
28 



There is, therefore, evidence suggesting that the Cape Flats population extends north 
to some locality on the Great Berg River and is there replaced by populations which 
are mainly pinkish-rufous, the latter not showing any appreciable dimensional 
difference nor difference in breeding-period. 

Lambert's Bay 

Three specimens, an adult and two juveniles, from Lambert's Bay, about 130 
miles north of Cape Town, are in the Transvaal Museum. The adult is slightly more 
colourful, about OOS/10/4, than the Saldanha Bay birds, and similar specimens 



326 



TAXONOMY OF THE KARROO AND 



collected by Roberts from near Port Nolloth, which he associated with his C. a. 
saldanhae: its dimensions are, wing 92, tail 71, bill 19. The juveniles taken in October 
are more richly coloured, about OOS/10/5, and compare with inland specimens from 
Klaver and Springbok areas. The material is too scanty on which to base any con- 
clusions, but it seems that 'greyish' birds somewhat similar to those at Saldanha Bay 
do occur on the coast at this point, and may connect with similar populations which 
have been found farther north at Port Nolloth. 



Swellendam and Deelfontein 

Before going inland, across the mountains, to Klaver, Van Rhynsdorp, and north 
to the Springbok area, specimens from east and north-east of Cape Town may be 
examined. 

A Layard specimen from Swellendam, which is about 150 miles due east of Cape 
Town and 30 miles inland, is snuff-brown, about OOS/6/5. It is in very fresh plumage, 
but is neither dated nor sexed: dimensions are, wing 83, tail 62, bill 18, first primary 
30. A rather lighter tone of colour, about OOS/7/5, is found in two adult specimens 
from Deelfontein, which is about 280 miles to the north-east of Cape Town and 
about 25 miles from De Aar where Roberts (1936a: 258) located his C. a. kurruensis, 
which was described as 'a very dark race'. One juvenile from the same locality, with 
wings and tail half-grown, is rather richer in colour, about OOS/6/7, while a second 
juvenile is noticeably browner, about OOS/9/5, and in this respect is very like birds 
in juvenile plumage from the Springbok area. The two adults were in fresh post- 
breeding plumage when taken in late February, which fits in more or less to the same 
breeding-cycle of the juvenile of the same colour taken in early March. The browner 
juvenile was taken in late January, and from the worn condition of its plumage had 
been in this dress for about 2-3 months. Dimensions are : 



Sex 


No. 


Wing 


Tail 


Bill 


F.P. 


$ 


1 


94 


69 


? 


28 


9 


1 


84 


62 


16 


26 


J- {1 


1 
1 


92 

? 


66 

? 


15 

? 


29 
? 



Apart from one specimen which does not fit in colour, nor apparently in breeding- 
cycle, the Swellendam and Deelfontein specimens, in colour at least, are similar. 
They are distinctly different from those in the areas so far examined, but appear to 
be not unlike birds in the Springbok area. 

Traka and Nels Poort 

Traka and Nels Poort are about 60 miles north and south of each other, and about 
250 miles east of Cape Town. Single Layard specimens from each of these localities 
are very similar in colour, but are rather duller brown than the Swellendam-Deel- 
fontein specimens. In the Traka and Nels Poort birds the lighter outer area of the 
feathers is almost entirely replaced by the darker centre which is dark brown, 
about OOS/6/4. The thin margin of lighter colour is about OOS/9/5, and the juvenile 



RED-BACK LARKS OF WESTERN SOUTH AFRICA 



327 



has almost pure white tips and margins. They are neither sexed nor dated. Dimen- 
sions are: 



Sex 


No. 


Wing 


Tail 


Bill 


F.P. 


? 
? 


1 

1 


92 
9i 


64 
68 


17 

? 


26 (Nels Poort) 
34 (Traka) 



Namaqua-Bushman-land Region 

Specimens from Klaver, on the Oliphant's River, and van Rhynsdorp, east to 
Calvinia and Carnarvon, and north to Kamieskroon, Grootberg, 1 Springbok, and 
the majority of those obtained near Port Nolloth are very alike in general colour, 
and will be considered together. A series of 63 specimens, largely from the Springbok 
area, have been examined. On the whole they seem to be a shade lighter than the 
Swellenden and Deelfontein specimens, being about OOS/8/5 as against OOS/6/5 ; 
the outer margins of the feathers are lighter, almost whitish in fresh specimens — 
particularly on the wing converts — and the dark central streaks are rather narrower 
and less diffuse. Below, the general colour is whiter, lacking much of the creamy- 
buff wash, which is particularly evident on the breast. 

A series of adults taken at different times of the year provides information on 
seasonal changes. Three males collected by Gill at Kamieskroon in September had 
enlarged gonads; they are very worn, and were almost certainly breeding. Speci- 
mens collected by us in the same locality in early December are just beginning post- 
breeding moult ; others from Klipfontein later in the month are just completing moult. 
Moult apparently begins on forehead and throat and works backwards towards the 
tail. Wing coverts are moulted early, but wing and tail moult begins later and in 
the usual manner, namely, starting at the junction of primaries and secondaries 
in the wing and proceeding away from that position, and commencing with the central 
pair of tail feathers. Specimens sometimes are in very fresh body plumage when 
wings and tail are barely half moulted. New feathers on the upper parts have a 
pinkish 'bloom' and have a very pale, nearly white edge on wing coverts, scapulars, 
and inner secondaries. During the year feathers become very much abraided, the 
pinkish 'bloom' disappears, pale edges disappear, and inner secondaries and central 
tail feathers in particular become very much worn. 

Young birds in juvenile plumage were more in evidence at Kamieskroon and 
Springbok early in December when adults were beginning to moult. None was 
found at Klipfontein later in the month when adults had nearly completed moult. 
(Young birds in this area were not specially sought.) The main feature of the juvenile 
plumage is that the dark centres of the feathers are much narrower, sometimes 

1 An Andersson locality. Wallis (1936: 285), in his biography of Andersson, notes that 'on July 2nd 
(1862) Andersson came to the ford of the Orange River (Sendelings Drift)'. After sending his wife and 
child ahead in the waggon 'he was able to rejoin them about the middle of July near the Buffalo River 
in Cape Colony'. He must therefore have been a good way south of Grootberg on the Orange River 
by the 29th July, the day on which the specimens were collected. The Buffalo River is presumably the 
one now named on maps as Buffel's River, which rises in the mountains south of Springbok. On some 
large-scale maps there is a locality marked Grootberg, about 13 miles south-west of Kamieskroon, and 
it is almost certain that this is the locality in which the specimens were collected. 

zoo. i, 11 xx 



328 



TAXONOMY OF THE KARROO AND 



almost completely absent on the body feathers of the back, but when present appear 
as black flecks rather than streaks. On the innermost secondaries the dark streaks 
are very little wider than the shaft, and in the central tail feathers it is only about 
half as wide as in adults. Whitish tips to body feathers are broad and produce a 
speckly appearance. The pinkish 'bloom' found in adults is lacking and because the 
dark feather centres are smaller the general effect is of a lighter and browner bird, 
although in fact the tone is very little different, about OOS/9/6. 

Seasonal change, mainly due to abrasion, is evident ; the worn plumage is appre- 
ciably duller. Four specimens, however, from widely scattered localities are rather 
lighter and browner in tone, about midway between true juvenile and adult. This 
difference may indicate a developmental variation, a distinctive first adult dress 
for example, or it may be an index of individual variation. 

Samples of population from the coastal plains around Port Nolloth show a high 
percentage of 'grey' phase birds. Roberts made a very interesting sectional picture 
of the population in this area when he collected between Klipfontein and Port 
Nolloth. Although his data suggests that 'grey' birds were limited to a narrow coastal 
belt about 25 miles in width (see Table below) , there are three specimens of 'red' birds 
in the National Collection which were obtained at Port Nolloth: two by Charles 
Reid in 1902 and one by C. H. B. Grant in 1903. Also out of six specimens collected 
by Meinertzhagen on the Klipfontein escarpment, about 45 miles inland, one is a 
'grey' bird. Meinertzhagen's specimen is the farthest inland record of the 'grey' 
phase. 



No. 


Locality 


Date 


Colour 


766 


Klipfontein 


17-8-37 


\ 


798 


25 miles east of P.N. 


19.8.37 






800 


„ 


M 




'Red' 


801 


>t »> 








802 


>» »» 


)( 


J 




808 


Port Nolloth 


1 1 


\ 


809 


,, 


,, 






811 


,, 


,, 




'Grey' 


822 


,, 


20.8.37 






841* 


25 miles east of P.N. 


tt 


) 




836 

840 


" 


21.8.37 
22.8.37 


} 'Red' 



* Date suggests that this number should be 831. 

Dimensions of 63 specimens from this region, including 'grey' birds from the coastal 
plains, are as follows : 



Sex 


No. 


Wing 


Tail 


Bill 


F.P. 


3 


38 


86-100 


62-74 


17-20 


28-36 


? 


21 


84-93 


58-67 


16-20 


26-32 


Juv. (| 


2 


89-91 
86 


70-74 
64 


17 

18 


28-32 
32 


?* 


1 


90 


65 


19 


28 



* Smith specimen: co-type of his Alauda lagepa. 



RED-BACK LARKS OF WESTERN SOUTH AFRICA 329 

Birds from this region have a higher degree of chromaticity than those in the Cape, 
Berg River, and Lambert's Bay areas; they are nearest to the Swellendam and 
Deelfontein specimens, but a shade lighter in tone, with a slight narrowing of the 
dark feather centres and a purer white on under parts. There is an obvious difference 
between juvenile and adult plumages due mainly to a great reduction in the width 
of the dark feather centres in juvenile feathers and the lack of pinkish 'bloom', typical 
of adults in fresh plumage. There is an indication that there may be an intermediate 
plumage between juvenile and full adult, and that coastal populations in the Port 
Nolloth area consist predominantly of 'grey' phase birds. These latter are somewhat 
similar in appearance to those found farther south at Lambert's Bay and Saldanha 
Bay. There is a large enough sample to show that females are slightly smaller than 
males and that the smaller samples from other areas fit into the range of measure- 
ments for this region. 

Orange River Mouth 

The Karroo Lark populations so far examined are readily linked together by 
similar dimensions and plumage pattern. (The significance of the colour differ- 
ence will be discussed later.) Immediately to the north-west of these populations, 
near the mouth of the Orange River, there are birds which appear to be geographical 
representatives of the Karroo Lark in which the dark streaks in the pattern are 
considerably reduced with a consequently higher proportion of the more brightly 
coloured areas (see Plate 37) . It is not known exactly where the change takes place 
and how it is effected. There is no obvious sudden change in the character of the 
country, which consists of monotonous rolling tracts of low scrub which gradually 
thin out to almost pure desert near the Orange River. The change was first observed 
in a specimen collected by the British Museum Expedition along the coast 38 miles 
north of Port Nolloth and a few miles south of the Orange River. The difference lies 
almost entirely in reduction in the width of the dark centres of the feathers. In the 
body feathers of the upper parts dark centres have almost completely disappeared, 
or are reduced to a thin and rather diffuse dark streak of the same general colour on 
the rest of the feather. Streaking on the breast is slightly reduced and is absent on 
flanks and belly. There is very little change in the pattern of wing and tail feathers. 

The specimen mentioned above, and six others collected at Grootderm, about 14 
miles up the Orange River, belong to the 'grey' group, while two others belong to 
the 'red' group. The 'red' tone is almost identical with that found in Springbok birds, 
about OOS/8/5, while the 'grey' tone is about OOS/9/4 and almost identical with 
Robert's 'grey' specimens from near Port Nolloth. Both the 'red' specimens were 
found in the company of 'grey' birds, and all are in the process of total moult. Three 
other specimens were found in the Transvaal Museum which had been collected in 
1942 at Orange Mouth. They are 'grey' birds, matching our specimens, except that 
they are stained with red soil or sand. Taken in September they must have been 
breeding. Dimensions of the twelve specimens are as follows : 



33° 



TAXONOMY OF THE KARROO AND 



Sex 


No. 


Wwg 


Tail 


Bill 


F.P. 




2 

6 
4 


86-89 

86-96* 

82-87 


63-67 
63-68 
61-64 


20-21 
20-21 
17-19 


28-30 
26-30 

26-2 7 



R. = 'red' phase. G. = 'grey' phase. 
* Next smallest are two specimens at 92. 

Important points to note about these Orange River specimens are that reduction 
in the amount of streaking is the only apparent difference from specimens in the 
Port Nolloth and Springbok areas, and that in the narrow coastal belt birds are pre- 
dominantly 'grey'. If the streaks were reduced to the point of disappearance the 
effect would be the appearance of the Red-back Larks which belong to regions north 
of the Orange River. Another connecting link between Karroo and Red-back larks 
is found in specimens from the next area to be considered. 

Witputs Area 

Witputs, a lonely police outpost, lies about 60 to 70 miles due north of Grootderm 
and at the southern tip of the Huib Plateau. A series of eighteen specimens was 
obtained there in late January. Environmental conditions were very similar to 
those in which the Karroo Lark lived in Little Namaqualand — namely, low sparse 
scrub mixed with prostrate succulents (see Plate 38a) . Although taken a month later 
than the Grootderm specimens the adults, of which there are thirteen, are only just 
completing moult. The adult plumage is darker and richer in colour than the 'red' 
Grootderm specimen, about OOS/7/6 (cinnamon-brown) as against OOS/8/5. 

Dark centres to body feathers of the upper parts have almost completely dis- 
appeared, although in occasional specimens — one out of the thirteen collected — the 
markings are rather more distinct and comparable with the Grootderm specimens 
(see Plate 37) . There is also an appreciable narrowing in the dark centres on the inner 
secondaries and central tail feathers. Little change is noticeable on the under parts 
except that the white is inclined to have a slight buffish tint. The worn breeding 
plumage is not represented. 

The five young birds are in moult and illustrate the transition from juvenile to 
first adult plumage. The moult appears to be a partial one, for there is no indication 
of wing and tail feathers being shed. Body moult proceeds in an anterior-posterior 
sequence. The difference in colour-tone between the two plumage phases is very 
apparent: the juvenile is a pale brown, about OOS/9/5, contrasting strongly with the 
OOS/7/6 of the fresh adult feathers. Interesting features are the reduction in the 
width of the dark centres of the central tail feathers and the increase in the width of 
the pinkish margins on the inner webs of the inner wing feathers. Dimensions are 
as follows: 



Sex 


No. 


Wing 


Tail 


Bill 


F.P. 


Ad.{f 
Juv. (f 


6 

7 

2 

3 


92-96 

83-87 
89-91 

78-84 


69-71 
59-64 
65-72 
61-65 


20-21 
17-18 

20 

17 


29-32 
26-28 
26-29 
26-30 



RED-BACK LARKS OF WESTERN SOUTH AFRICA 



33i 



Although the Witputs birds are clearly different to those at Grootderm, they are 
obviously related, for the differences are only slight quantitative changes. 

Aus Area 

Aus is a small village about 80 miles north of Witputs, about 60 miles inland from 
Luderitz Bay, and on the Namib Desert side of the main backbone range of moun- 
tains. It is easily reached by road or rail and has been a collecting centre for a 
number of ornithologists. Red-back larks in this area have given rise to some con- 
troversy. 

Roberts (1937 : 88) reported that whilst delayed by a mechanical breakdown about 
30 miles north of Aus, on the Helmeringhausen road, he collected a series of larks 
which he claimed to be identical with Pseudammomanes erythrochlamys (Strickland) 
from Walvis Bay. Subsequently he camped 'in the flats below the hills to the west 
of Aus', where he obtained a further series of Red-back larks. These he considered 
were different from P. erythrochlamys 'in having the inner lining of the wing-quills 
with only a trace of pinkish on the inner edges — not broadly pinkish — and the middle 
tail feathers above with a broad central stripe of dark greyish-brown ; the outer tail 
feathers are also on the whole darker, only the outer web of the outermost being 
pinky-whitish, the rest of the feather blackish with only the tip pale; the outer 
webs of the primaries above is only very thinly or not at all pinkish'. He also gives 
the following dimensional differences : 



Sex 


No. 


Wing 


Tail 


Tarsus 


Culmen 


P. barlowi 1 ~ 
P. erythrochlamys \ ~ 


7 

2 

8 
5 


93-97 

83-87 
89-95 

84-85 


68-72 

61-5-64 

67-74 

64-69 


24-5-26 

24-5-26 

26-28 

24-26 


18-19 
16-16-5 
17-18-5 
14-5-16 



He considered that these were sufficient criteria for the recognition of a new species 
and gave them the name P. barlowi. 

Hoesch and Niethammer (1940: 60) visited Aus in late December and early 
January 1938-1939. Red-back larks were collected only on the farm of Kubub, 
which is about 2-3 miles south of Aus, except for one found about 30 miles farther 
west, at Tschaukaib, on the road to Luderitz Bay (see Fig. 1.) On the evidence of 
these specimens they disagreed with Roberts, stating that the dark stripe in the 
central tail feathers is variable, and put his P. barlowi into the synonymy of P. 
erythrochlamys. They do not record that they had either found or examined typical 
specimens of P. erythrochlamys. Roberts had this advantage, for there was a speci- 
men from the Kuiseb River in the Transvaal Museum collected in December 1928 
by R. D. Bradfield. 1 

It is almost certain that Roberts would have made his points clearer in the fuller 
account which he proposed to publish, for the important factor correlated with the 
difference he observed is the occurrence of large tracts of shifting sand-dunes. In 
his preliminary account the only clues he gives are that the birds 30 miles north of 

1 Incidently, Roberts collected two specimens at Rooibank, 30 miles up the Kuiseb River, in September 
1 94 1 : he did not have an opportunity to comment on them in print. 



332 



TAXONOMY OF THE KARROO AND 



Aus were 'found in large numbers on some dunes, but not in the plain near by', 
while of those west of Aus he merely states they were 'on sandy dunes'. Details of 
the area in the vicinity of Aus are shown on the accompanying sketch map. 

To the north of Aus the dry-bed of the Koichab River forms the southern boun- 
dary of an immense area of sand-dunes which stretches for about 200 miles northwards 
to Walvis Bay at an average depth of about 70 miles from the coast. The Helmering- 



2.6 













27 



IS 




-U-l — \- 



2S 



ri'les 



lb 



TlR^S 



ms 








AusteC 



27 



(6 



Fig. 1 



Details of Aus area showing southern limit of shifting sand-dunes and localities in which 
Red-back larks were collected. 

B.M. = B.M. Expedition collecting locality. 
R. = Roberts's collecting locality. 

hausen road, 30 miles north of Aus, touches an encroaching arm of the dunes. In 
fact it was evident that the road we used in 1950 is a new trace circumventing the 
dunes and that Roberts was almost certainly on the old road. 

We explored the dunes about 30 miles farther west than Roberts did. They lay 
across our path in the form of a pinkish-red barrier of fine wind-blown sand rising 
several hundred feet. On the margins of the dunes there were numerous scattered 
clumps of a spiky grass, Aristida sp. Red-back larks were found living in associa- 
tion with this grass. (Conditions were similar to those at Tsondab Mund shown on 
Plate 386.) 

South of the dunes the country consists of a jumble of granite hills and kopjies 
rising from plains of firmer sand and gravel covered with low scrub, varying in 
density according to local conditions. Red-back larks may be found almost any- 



RED-BACK LARKS OF WESTERN SOUTH AFRICA 



333 



where in this scrub. Niethammer records that he found them in small parties on 
the western edge of the Kubub plain where there were patches of small bushes. We 
came across them in several places north, south, and west of Aus. 

There is no doubt that Aus (scrub) birds are different. They are clearly inter- 
mediate between the Aus (dune) and the Witputs birds, which also inhabit a similar 
sort of scrub, but smaller and sparser, frequently with a high proportion of succu- 
lents, and growing on a redder type of sand. Twelve specimens collected by us near 
Aus in late January and early February are adults just completing moult and are in 
perfectly fresh plumage. They are much paler than the Witputs birds, about OOS/9/5 
as against OOS/6/6. Dark centres to body feathers on the upper parts are never 
blackish as they sometimes are in Witputs specimens, but are a darker tone of the 
same general colour. Dark streaks on the central tail feathers are on the whole 
appreciably reduced, and on the innermost secondaries are little more than the 
thickness of the shaft. Below, the streaking on the breast is much less distinct 
(see Plate 37). In Roberts's specimens, collected in July, the plumage is fairly worn, 
but the only change is in the loss of most of the pinkish 'bloom' which appears to be 
characteristic of all fresh adult plumages. There are no juveniles. Dimensions of 
British Museum and Transvaal Museum specimens are : 



Sex 


No. 


Wing 


Tail 


Bill 


F.P. 


3 


14 

6 


91-100 
86-92 


67-76 
63-71 


19-21 
18-19 


27-38 
29-32 



Namib Dunes 
North of Aus 

The Aus (scrub) birds are more like the sand-dune specimens from 30-40 miles 
north of Aus than the Witputs birds. Of six specimens collected on the dunes on 
the 31st of January, four are in the final stages of moult, while two are in half moult 
and still showing the appearance of the old worn plumage. The colour of the upper 
part in the fresh and worn plumage is indistinguishable from that of the Aus 
(scrub) birds, being about OOS/9/5 when fresh and about OOS/10/5 when faded. 
The main difference lies in the extreme narrowness of the dark centres of the central 
tail feathers which are reduced more or less to the width of the shaft. On the inner- 
most secondaries it has disappeared altogether, even the shaft being the same colour 
as the web. Differences on the under parts are more noticeable : the streaking on the 
breast is reduced very considerably, both in the number of feathers with dark centres 
and the width of the dark centre, which is not blackish, as in the population samples 
already examined, but about the same colour as the upper parts (see Plate 37) . Also 
the white of the under parts is washed with pale buff. In adults there is a greater 
amount of pinkish on the underside of the wing (on the inner margin of the inner 
web). It may be noted that this and other characters, such as the narrower width 
of dark centres on central tail feathers and inner secondaries, are associated with 
juvenile plumage in the Aus (scrub) and Witputs areas. 

In the preserved specimens the bills of the dune birds have dried off to a dark 



334 



TAXONOMY OF THE KARROO AND 



brownish-horn colour, whereas in the scrub birds they are blackish-horn. This 
difference is not reflected in the data noted in the field, but direct comparison of 
fresh specimens was not made and unless some standard colour nomenclature is used, 
field descriptions, even of the same colour, often vary from day to day. 



Sex 


No. 


Wing 


Tail 


Bill 


F.P. 


9 


8 
9 


91-97 
83-90 


68-71 
60-70 


18-20 
16-20 


28-32 
23-29 



Tsondab Mund 

A series of specimens was obtained at Tsondab Mund, about 180 miles north of 
Aus and 20 miles south of the Kuiseb River. The Tsondab is a seasonal river which 
has its origin in the Naukluft Mountains, and disappears in the sands of the Namib 
Desert, about 50 miles from the coast. It lies in a gravel plain flanked by mountain- 
ous dunes which eventually form a barrier across its course (see Plate 386). The dunes 
are very similar to those at Aus. A series of eighteen specimens was obtained on 
the 5th and 6th of March, of which one is in juvenile plumage. The adults are either 
in extremely worn breeding dress, with gonads large but apparently subsiding, or 
just beginning to moult. They are, therefore, about two months later in their 
breeding-cycle than the population near Aus. A few are fairly well advanced in 
moult, sufficient to show that the colour of the fresh plumage is exactly similar to the 
Aus (dune) birds, being about OOS/9/5 when fresh and fading to about OOS/10/5, 
On the underside also colours and markings are identical, but in the worn condition 
the breast stripe almost disappears (see Plate 37). The juvenile is similar to the worn 
adult in colour, but is distinguishable by pale tips to most of the feathers, giving it 
a speckled appearance. Dimensions are: 



Sex 


No. 


Wing 


Tail 


Bill 


F.P. 


3 

Juv. $ 


11 

6 


88-98 

83-88 

82 


62-70 

60-66 

60 


19-21 
18 
16 


25-33 

25-31 

29 



Walvis Bay 

The Kuiseb River forms the northern boundary of the main Namib sand-dune area. 
It is not certain exactly where C. J. Andersson obtained the specimens on which 
Strickland based his descriptions of Alauda erythrochlamys. Of those in the British 
Museum two carry the information 'sandy flood bed of the Kuiseb River'. He was 
stationed both at Walvis Bay and Sweppmansdorp (now Rooibank), about 30 miles 
up the Kuiseb. Roberts obtained specimens in 1941 at Rooibank 'at the edge of 
the sand dunes'. We spent a few days higher up, at Swartbank, but were on the 
north side and separated from the dunes by the Kuiseb in flood. 

Of six specimens in the British Museum, two adults are dated November. They 
are in very worn plumage and compare exactly with worn and faded specimens from 
Tsondab Mund, about OOS/10/5. Of two in juvenile plumage one is dated May, 
and judging by the extent of wearing compared with a March specimen from Tsondab 



RED-BACK LARKS OF WESTERN SOUTH AFRICA 



335 



Mund, must have been hatched out about the same month. It seems, therefore, that 
the breeding-cycles along the Kuiseb and Tsondab coincide. Dimensions are: 



Sex 


No. 


Wing 


Tail 


Bill 


F.P. 


? 
Juv. £ 

p 


4 

i 
i 
3 


90-93 

89 

87 
89-90 


67-68 
62 

•86 
67-68 


19-21 

17 

18 

17-19 


26-28 
30 

28-29 



Roberts and Meinertzhagen quote Swakopmund in the distribution of this lark. 
So far as I know specimens have not been found in that locality. Although there 
are dunes near the coast at Swakopmund we found them entirely lacking both in 
Aristida grass and Red-back larks. Roberts also mentions that Andersson took eggs 
of this species at Otjimbinque, and these are in the British Museum. As the eggs 
are not accompanied by the birds they belonged to their proper identification will 
remain uncertain until they can be compared with eggs known with certainty to 
belong to A. erythrochlamys. 

DISCUSSION 
Dimensions 

The localities mentioned above are indicated on the accompanying map (Plate 36) . 
The sample of specimens from each locality is, in the majority of instances, too small 
in relation to the number of factors affecting dimensions, such as, for example, the 
amount of wear, to obtain accurate statistical figures, but the measurements taken 
are approximate enough to give a general picture of correlation between the various 
populations. The only factor which need be taken into account is sex, but even 
here a high degree of accuracy in sexing cannot be claimed. Juvenile measurements 
do not appear to vary noticeably from those of adults. In any case, if they were to 
be dealt with separately, account would have to be taken of the first adult com- 
pound plumage in which juvenile wing and tail feathers are retained, but this stage 
does not appear to be recognizable. Measurements of males and females, including 
the juveniles of each sex, from the various localities, are compared in Tables 1 and 2, 

Table i. Summary of the Dimensions of Males of All the Populations of Karroo 

and Red-back Larks 



Locality 


No. 


Wing 


Tail 


Bill 


F.P. 


Cape Flats ....... 


5 


88-94 


62-68 


18-20 


28-32 


Berg River .... 








3 


9i 


62-68 


19-20 


31-32 


Lambert's Bay .... 








1 


92 


7i 


19 


— 


Swellendam .... 








2 


92-94 


66-69 


15 


28-29 


Namaqua-Bushman-land Region 








40 


86-100 


62-74 


17-20 


28-36 


Orange River .... 








8 


86-96 


63-68 


20-21 


26-30 


Witputs ..... 








8 


89-96 


65-72 


20-21 


26-32 


Aus (scrub) .... 








14 


91-100 


67-76 


19-21 


27-33 


Namib: Aus (dunes) . 








8 


91-97 


68-71 


18-20 


28-32 


Tsondab Mund .... 








11 


88-98 


62-70 


19-21 


25-33 


Walvis Bay .... 








5 


87-93 


67-68 


12-21 


26-28 


Total .... 








105 


86-100 


62-76 


17-21 


25-36 



ZOO. I, II 



Yy 



336 



TAXONOMY OF THE KARROO AND RED-BACK LARKS 



Table 2. Summary of the Dimensions of Females of All the Populations of Karroo 

and Red-back Larks 



Locality 


No. 


Wing 


Tail 


Bill 


F.P. 


Cape Flats .... 
Berg River .... 
Swellendam .... 
Namaqua-Bushman-land Region 
Orange River .... 
Witputs ..... 
Aus (scrub) . . ■ 
Namib: Aus (dunes) . 
Tsondab Mund .... 
Walvis Bay .... 








4 
1 

1 
22 

4 
10 
6 
9 
7 
1 


82-88 
88 
84 

84-93 

82-87 

78-87 
86-92 
83-90 
82-88 
89 


56-61 

61 

62 
58-67 
61-64 
59-65 
63-71 
60-70 
60-66 

62 


18-20 

20 

16 
16-20 
17-19 
17-18 
18-19 
16-20 
16-18 

17 


27-33 

27 

26 
26-32 
26-27 
26-30 
29-32 
23-29 
25-31 

30 


Total .... 








65 


78-93 


56-71 


16-20 


23-33 



which summarize the full data given in the Tables of Measurements' at the end. 
From this it is seen that none of the samples, or group of samples, under examina- 
tion is readily separated from the others. 

The uniformity apparent in this general picture can be tested by examination of 
the frequencies of the dimensions taken. It seems legitimate to emphasize any small 
differences by combining lengths of wing, tail, and bill (first primary measurements 
being omitted only because they are less complete) . This has been done separately 
for males and females and the result is shown logarithmically in Fig. 2. Males have 
a mean value of 180 mm. and females 167-5, with standard deviations of 6 and 5 
respectively. The main point, however, is that the distributions are approximately 
symmetrical, suggesting that the sample belongs to a more or less uniform population. 

In fact the sample is so uniform that a cline is not even evident. When Namib 
dune populations are tested against populations from the Namaqua-Bushman-land 
area an almost exact correlation is obtained. For instance, in Fig. 3 the size-fre- 
quencies of the wing lengths of the males of these two populations are shown as 
histograms and are plotted logarithmically. The difference of the means (o-i mm.) 
is so low that it is clearly of no significance; about 95 per cent, of random samples 
of a single population would show a mean difference of this magnitude, or greater. 

At this point it may be noted that Roberts (1937 : 97) distinguished his Calendu- 
lauda guttata calviniensis only on size ; he gives wing measurements as <$ 99, ? 90 
(by my measurements 98 and 89), both of which lie within the range given here. 
Apparently he only had two specimens and a larger sample from the same locality 
obtained by Meinertzhagen shows a wider range: 5 males, wing 90-96, 2 females, 
wing 85-89. In this short series the mean wing length of the males, 93-9 mm., differs 
from the mean length of the Namaqua-Bushman-land populations by only 1 mm., 
and the type male alone differs from the mean by an amount greater than the 
standard deviation of the populations. 

Colour and pattern 

In general appearance birds are coloured above in various tones of reddish-orange 
which is either plain or marked with very dark streaks: the under parts are whitish, 



2 I 5 



>.o» o»i 




Fig. 2. Size-frequencies of combined wing, tail, and bill lengths of 97 males and 64 females from 
all the populations of Karroo and Red-back larks. 



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37 specimens; circles and unbroken line. Mean 92-9 mm. ; a, 3-0. (b) Namib dune populations, 

23 specimens; crosses and broken line. Mean 92-8 mm. ; a, 2-7. 



338 



TAXONOMY OF THE KARROO AND 



with dark streaks at least on the breast. In all body-feathers, both above and below, 
the basal portion (at least two-thirds or more) is uniformly very dark grey. The 
usual pattern of the exposed tip consists of a dark central streak, flanked by a 
coloured area of medium tone, and a thin pale outer margin (see Fig. 4). On the 
whole these parts are fairly sharply defined. The pale outer margin is most evident 
in the juvenile plumage, where it forms a fairly broad whitish tip. Pale margins 
are also present sometimes in new adult feathers, but they are usually very narrow 




Concealed 



Fig. 4. Diagrammatic representation of the colour pattern of a feather 
taken from the centre of the back. 



and soon wear off. In some localities, particularly the Springbok area, broad whitish 
margins to the feathers of the wing coverts contrast sharply with blackish centres 
and form a pattern which persists throughout the season. 

The general colour of the upper parts is mainly determined by the coloured area 
of medium tone. It is this colour which has been identified and used for comparative 
purposes. In all the specimens examined the colours of this area belong to the same 
hue, as identified by the Colour Atlas. They vary only in the degrees of lightness 
and chromaticity. An attempt has been made to show the extent of this variation 
in Fig. 5. Two general points of special interest which are illustrated are that the 
less colourful and rather paler forms are distributed along the coast from Cape Town 
to the mouth of the Orange River ; otherwise birds of the main population have the 
same chromatic value, varying only in degree of lightness. 

As well as variation in tone the coloured area varies in the amount of the exposed 
tip it occupies. In the northern Namib birds it occupies the whole of the exposed 
tip, the dark centres being non-existent except in a few of the breast feathers. In 
Aus birds the dark centres are slightly more pronounced, and become more so at 



RED-BACK LARKS OF WESTERN SOUTH AFRICA 



339 



Witputs and the Orange River. In the Springbok region there is a sudden increase 
in the ratio of dark centres to coloured area, so much so that in this particular feature 
birds at Orange River mouth are more unlike birds from Port Nolloth, only 60 miles 



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LIGHTNESS VALUES 



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Fig 



. 5. Graphical representation of colour variation based on Villalobos's Colour Atlas. The 
area ringed indicates the grey coastal population. 

Aus ; B.R. = Berg River ; C = Cape ; D = Deelfontein ; N = Namib ; N.P. = Nels Poort ; O = Orange River ; 
Springbok ; SW = Swellendam ; S.B. = Saldanha Bay ; T = Traka ; W - Witputs ; (G) = Grey phase ; (R) = Red 

phase ; (J) = Juvenile. 



away, than birds from the Namib dunes. Birds from Springbok southwards could 
be described as being heavily streaked, with a tendency in some places in the south 
for the dark centres to occupy nearly the whole area of exposed feathers. Birds 
north of Springbok are lightly streaked, with a tendency in the north for the coloured 
area to occupy nearly the whole of the exposed part of the feather. There is there- 
fore a more or less graded change in pattern correlated with north and south 



340 TAXONOMY OF THE KARROO AND 

distribution. There does not seem to be any correlation between colour and pattern. 
The populations which have similar broad streaks are variable in colour. 

Developmental stages 

The juvenile plumage is distinct from the adult plumage. Most of the feathers 
have a fairly broad whitish margin at the tip, the dark central streaks are narrower, 
and in general appearance the plumage lacks the particular pinkish bloom which 
seems to be typical of all adult fresh plumages. On the whole the colour of juvenile 
plumages most closely resembles worn and faded adult plumages, but it may be 
noted that of two young birds obtained in the Deelfontein area, by the same collector 
in 1901, a very young March specimen is more richly coloured than adults, while a 
rather worn January specimen is rather lighter and browner and agreeing with the 
more usual relationship between juvenile and adult. The significance of this differ- 
ence does not seem to be apparent at this stage. Moult from juvenile to first adult 
plumage seems to take place slightly later than the adult post-breeding moult. Only 
the body-feathers are moulted and therefore the first adult plumage is compound. 
It may be noted that it is possible that this stage may be rather less deeply coloured 
in populations of deeply coloured adults. Moult proceeds in an anterior-posterior 
sequence. 

In the adult plumage there is an appreciable seasonal change due to wear and 
fading, new feathers having a noticeable pinkish bloom. In the material studied 
there is no indication of pre-nuptial moult, partial or otherwise, so that the moult 
cycle seems to be Juvenile — > Compound Annual — > Simple Annual (repeating) . In 
the adult post-breeding moult there is a complete change of feathers. Body moult 
proceeds in the same sequence as in the juvenile and is usually rather ahead of 
wing and tail, in which the sequence follows the normal pattern, beginning at the 
junction of primaries and secondaries in the wing and the central pair of feathers 
in the tail. 

The pattern of development therefore is uniform throughout the group, and in 
the rather limited material examined there seems to be rather less variation in the 
juvenile than in the adult, the former being constantly browner in colour because of 
the narrower dark feather streaks. 

Breeding-cycle 

In the northern sector of the range there are indications of a progressively later 
breeding-cycle correlated with decrease in latitude. Birds at Kamieskroon (30 south) 
had commenced post-nuptial moult in late November, whereas at Tsondab Mund 
(24 south) it was early March before birds reached the same stage. In the southern 
sector, however, the correlation is confused, as might be expected when the distri- 
bution spreads eastwards. The factors determining the breeding-cycle have not been 
investigated. A similar gradation in breeding-cycle was found in the Long-bill lark, 
Certhilauda curvirostris (see Macdonald, 1952). 

Habits 

Regarding the habits of these birds, Smith (1843) stated of his A. codea that 'when 



RED-BACK LARKS OF WESTERN SOUTH AFRICA 



34i 



disturbed they fly to a distance, and then perch upon the summit of some dwarf 
shrub' ; and of his A. lagepa 'on descending from its aerial flight commonly perches 
on the shrub nearest to the point where it descends'. Notes by Andersson (1872), 
according to Gurney, are not related with certainty to any birds belonging to this 
group. Layard (1867 : 209) under A . codea records some observations on the habits of 
nesting birds. Grant, in Sclater (1911: 256), records (under Mirafra nivosa — an in- 
valid name, see footnote p. 321) that 'it frequents open flats and the tops of mountain 
ranges, and is usually in pairs. The call is a whistle, and the bird is fond of perch- 
ing on the tops of low bushes and scrub, especially if disturbed'. Jack Vincent found 
his birds at Blaauwberg beach 'among low scrub on sand dunes'. An account of the 
nesting and other habits of the Cape race is given by A. W. Vincent (1946: 466). 
Roberts makes a few references to habits. Hoesch & Niethammer (1940: 224) 
recorded of the Red-back lark that they found it in small parties on the western 
edge of the Kubub (Aus) plain where there were patches of small bushes, and also 
that they run very fast over the sand and stop motionless in the cover of bushes. 

Of birds in the Springbok area we noted that they were usually found in places 
where scrub became sparse and stunted and the soil loose and sandy or gravelly. 
On the ground they ran vigorously. They were never seen to make more than short 
low flights which were rather weak and fluttering. In our experience they always 
landed on the ground, making what appeared to be a 'pancake' landing, but they 
would jump on to low scrub. Local populations seemed to be thinly scattered in ones 
and twos. At Grootderm, near the mouth of the Orange River, they were found in 
rather more arid conditions, in depressions between low hills on soft sand with 
scattered stones and rocky outcrops, and practically no scrub, but what scrub there 
was the birds used for cover, running from one tuft to another. On the few occasions 
on which observations were made none were seen to perch. At Witputs they were 
again associated with open scrub, about 18 inches high, and soft sand. They were 
relatively plentiful, usually in scattered pairs, but took a good deal of finding until 
one became acquainted with the places they liked and their habit of creeping about, 
mouselike, of standing motionless under a clump of scrub, or of running at great 
speed with head down, but then frequently giving themselves away by jumping up 
on top of the scrub. They were difficult to flush, and flew low for very short 
distances. At Aus the picture was much the same : birds were most frequently found 
on sandy patches with sparse stunted scrub, and when disturbed ran for long dis- 
tances. Sand-dune birds from north of Aus and Tsondab Mund were not noticeably 
different. They lived in close association with Aristida grass, sheltering in the clumps, 
or running with amazing speed from one clump to another, easily outpacing our 
clumsy efforts to catch up with them. Probably because of this we noted that they 
seemed to be wilder than other Red-backs, but then they were living in a more 
exposed environment. They sometimes jumped up on top of the grass tufts, but 
were never seen to alight there from flight. A display flight was noted, in some ways 
reminiscent of the skylarks. Birds climbed up to 100 feet or so then fluttered hori- 
zontally for a short distance uttering a rather musical note which was written down 
as 'chek-chek-chek-chek-tae' : they would drop 10 or 20 feet, flutter again for a short 
distance, then drop suddenly to earth, and run. A variation of this note was 



342 TAXONOMY OF THE KARROO AND 

sometimes uttered by a bird standing in or on a clump of grass : it was recorded as 
'tchee-tchee-tchee-chr-r-r-r'. 

It seems clear, therefore, that the birds known as Karroo and Red-back larks live 
in similar habitats, namely, areas of soft sand where vegetation is sparse, and that 
there seems to be a good deal of similarity in their habits. 

Nomenclature 

It would be convenient to summarize the foregoing and leave the matter there, 
for it is obvious that much more data has to be obtained before an accurate picture 
of the taxonomy of this group can be presented. But for cataloguing purposes the 
question of nomenclature has to be dealt with. 

The general picture is of a group of populations which are practically identical 
dimensionally, but variable in several other characteristics, particularly colour-tone 
and feather pattern. These variations appear to be independent of each other, but, 
for the most part, they can be correlated with distribution. Streaking is heaviest 
in the southern birds and almost completely disappears in the northern. Colour 
bears a broad general relationship to soil colour. In fact, putting both together, 
plumage colour and pattern bears some relation to soil colour and pattern. In the 
smooth fine sands of the Namib dunes birds are plain; whereas in the Springbok 
area, for example, where there is much more gravel and stones and prostrate vege- 
tation breaking up the surface of the sandy localities frequented by these birds, they 
are patterned. A striking colour variation is associated with a narrow coastal belt 
which has greyish sands and frequent coastal fogs. 

It seems highly probable that these variable characteristics are largely phenotypic 
in origin and of less significance taxonomically than those which are less variable, 
such as dimensions; and therefore, taking into consideration the allopatric nature 
of the distribution, it can be concluded that all these various populations are repre- 
sentatives of a single species. 

This lark seems to be related to the Long-bill lark, Certhilauda curvirostris, which 
occupies rather stonier types of country within practically the same area of distribu- 
tion, and is the type species of the genus Certhilauda. The specific name of the Karroo 
and Red-back larks therefore is Certhilauda albescens (Laf resnaye) . 

The question as to which populations should carry distinctive names is more 
difficult to answer. Grey birds appear to be restricted to coastal localities between the 
Cape and the Orange River. Cape area populations seem to be entirely grey and of 
a distinctive tone and therefore may be given racial status. But from Lambert's 
Bay north to Port Nolloth populations are mixed grey and red, though predominantly 
the former. This mixing seems to have been the main reason for Roberts's recogni- 
tion of two species and Meinertzhagen's reversion to a single polymorphic Karroo 
Lark not divisible into races. My own opinion is that the coastal strip in which 
grey birds predominate is no wider than the zone of overlap or infiltration which one 
would expect to find between two geographical forms whose differences are not 
apparently intergraded (see map, Plate 36). Some overlap may be due to birds 
wandering in the non-breeding season and the balance of grey over red being main- 
tained no doubt by those factors which are the primary cause of a grey phase being 






RED-BACK LARKS OF WESTERN SOUTH AFRICA 343 

established in this area. I think, therefore, that it presents a better picture of the 
taxonomy of the species to give them racial status and, until more is known about 
them, to tack them on to the Saldanha Bay form. 

The inland red populations are, in my opinion, not well enough known as yet, 
throughout their wide distribution, to be regarded as anything other than a single 
race, guttata. In the north the scrub and dune populations are distinct and have 
been named, but in the Witputs area, and near the mouth of the Orange River, there 
are distinct populations for which names have to be provided. Regarding the latter, 
the type is one of a series of grey birds, and it may well be that when more becomes 
known about populations in this area that an inland red form will be distinguished. 

The nomenclatorial picture, therefore, is as follows: 

Certhilauda albescens (Lafresnaye) 

(1) C. a. albescens (Lafr.) 

Alauda albescens Lafresnaye, Rev. Zool. 1839: 259: Blaauwberg Beach, Table 
Bay. 

Alauda codea Smith, Zool. of S. Africa, 1843, pi. 8j: Karroo plains, between 
the Oliphant's and Orange Rivers. (Probably Cape Flats.) 

Characteristics. General appearance of upper parts light drab, broadly streaked 
with sepia. Below whitish broadly streaked with sepia on breast, lightly streaked 
on flanks and almost entirely without streaks on belly. 

Distribution. Cape Flats, as far north as Berg River. 

(2) C. a. saldanhae (Roberts) 

Calendulauda albescens saldanhae Roberts, Ann. Trans. Mus. 1936: 258: 
Saldanha Bay, Cape Province. 

Characteristics. Similar to previous race, but with a pinkish-rufous wash on 
the upper parts. The extent to which this feature is constant in the areas indi- 
cated is not certain. 

Distribution. Saldanha Bay, Berg River, Lambert's Bay, and northwards along 
the coast to Port Nolloth. 

(3) C. a. guttata (Lafr.) 

Alauda guttata Lairesnsye, Rev. Zool. 1839: 259: Elephant's (Oliphant's) River, 
Cape Province. 

Alauda legepa Smith, Zool. of S. Africa, 1843, pi. 87: between the Berg and 
Orange Rivers. 

Calendulauda albescens karruensis Roberts, Ann. Trans. Mus. 1936: 258: de 
Aar, Cape Province. 

Calendulauda guttata calviniensis Roberts, Ostrich, 1937 (97) : Calvinia, Cape 
Province. 

Characteristics. Similar to previous races in the extent of sepia streaking above 
and below, but general colour of upper parts about snuff -brown to Mikado-brown, 
zoo. 1, II zz 



344 TAXONOMY OF THE KARROO AND 

Some variation in colour is evident, but so far not clearly associated with 
distribution. 

Distribution. Inland areas from Swellendam in the south, north-east to de 
Aar and west to Springbok, and apparently sometimes reaching the coast, as at 
Port Nolloth and Lambert's Bay. 

(4) C. a. patae new race 

Characteristics. Upper parts very lightly streaked and dark central streak on 
inner secondaries and central tail feathers much reduced. Below, streaks con- 
fined entirely to breast. Two colour phases known from the type locality, one 
similar to the general colour of C. a. guttata and the other to the general colour of 
C. a. saldanhae. 

Distribution. South bank of the Orange River near its mouth to coast about 
10 miles south. 

Type. One of the 'grey' phase. Male ; collected at Grootderm, Orange River, 
Little Namaqualand, lat. 28 31' S., long. 16 38' E., alt. 500 ft., on 17th 
December 1949 by the British Museum South West Africa Expedition (1949- 
1950). Register number 1950:50:936. Wing 89, tail 67, bill 21. Iris brown, legs 
pale yellow-grey, bill black. Approaching final stages of moult. 

Remarks. The series consists of seven 'grey' phase and two 'red' phase, 
obtained in mid-December: three 'grey' specimens in the Transvaal Museum 
were obtained at Orange Mouth in September. 

(5) C. a. cavei new race 

Characteristics. On the whole rather less streaked above than the Orange 
River birds, and upper parts darker and richer in colour than 'red' specimens 
from that area, about cinnamon-brown. Streaking below much the same. 

Distribution. At the southern end of the Huib Plateau in the vicinity of 
Witputs, Great Namaqualand. 

Type. Male; collected 5 miles south-west of Witputs, Great Namaqualand; 
lat. 27 35' S., long. 16 42' E., alt. 4,000 ft. ; on 26th January 1950, by the 
British Museum South West Africa Expedition (1949-1950). Register number 
1950:50:922. Length of wing 96, tail 71, bill 21. Iris dull brown, legs pale grey, 
bill dark grey. 

Remarks. Eighteen specimens, three of which are juvenile, were obtained in 
late January. 

(6) C. a. barlowi (Roberts) 

Pseudammomanes barlowi Roberts, Ostrich, 1937: 95: 8 miles west of Aus, 
Great Namaqualand. 

Characteristics. Plain above, with no indication of dark central streaks on body 
feathers : streaks on inner secondaries and central tail feathers reduced to little 
more than a thin line. General tone of colour much lighter than Witputs birds, 



RED-BACK LARKS OF WESTERN SOUTH AFRICA 345 

about sayal-brown. Below, dark streaks on breast distinctly lighter, narrower, 
and occupying a smaller area: white ground colour washed with pale buff. Bill, 
blackish-horn. 

Distribution. Vicinity of Aus, Great Namaqualand: extending at least 20 miles 
west, 6 miles north, and about 3 miles south. 

(7) C. a. erythrochlamys (Strickland) 

Alaudaerythrochlamys Strickland, Contr.Orn. 1852: 181: Damaraland (probably 
Kuiseb River near Walvis Bay). 

Characteristics. Plain above and similar in colour-tone to previous race, but 
lacking dark centres to inner secondaries and central tail feathers. Below, 
streaking on breast much reduced, and dark cinnamon-brown rather than sepia ; 
white ground colour more washed with buffish ; and more pinkish-buff on under- 
side of wing. (It may be noted that several of these features are found in the 
juvenile stages of C. a. barlowi and C. a. cavei.) Bill, brownish-horn. 

Distribution. Namib sand-dune area from the Koichab River basin just north 
of the Aus, north to the Kuiseb River as far as Walvis Bay, and inland as far as 
the dunes extend and apparently where spiky Aristida grass occurs. 

SUMMARY 

i. Samples of various populations of Karroo and Red-back larks have been 
examined, particularly as regards dimensions, colour, pattern, moult, breeding- 
cycle, development, habits, and habitat. 

2. The differences found are no greater than might be expected in random samples 
of the same species, and it is concluded that Karroo and Red-back larks can be 
regarded as one species, Certhilauda albescens (Lafresnaye) . 

3. Variation in colour and pattern are broadly correlated with the colour of the 
soil and the pattern of the environment ; plainness being associated with an environ- 
ment of smooth sand and streakiness with a broken pattern. 

4. Although colour and pattern vary independently, seven geographical races 
based on these characteristics can be recognized. Two are newly described. 

5. It is noted that data is still very inadequate and much still remains to be found 
out, particularly as regards distribution and variation in populations in Cape 
Province, and that the racial picture presented for birds in that area may be subject 
to amendment. 

REFERENCES 

Andersson, C. J. 1872. The Birds of Damaraland. xlviii+394 pp., 3 figs. London. 
Ayres, T. 1874. Additional list of and notes on birds obtained in the Republic of Transvaal. 

Ibis, 1874: 101-107. 
Harding, J. P. 1949. The use of Probability Paper for the graphical analysis of poylmodal 

frequency distribution. /. Mar. biol. Ass. U.K., 28: 141-153. 
Hoesch, W., & Niethammer, G. 1940. Die Vogelwelt Deutsch-Siidwestafrikas. /. Orn., 

Lpz. Supplement, viii+404 pp., 8 pis., 76 text-figs. 



34 6 



TAXONOMY OF THE KARROO AND 



Lafresnaye, Fr. de. 1839. Quelques nouvelles especes d'oiseaux. Rev. Zool. 1839: 257-259. 
Layard, E. L. 1867. Birds of South Africa. . . . xvi + 382 + xxipp., 1 pi. Cape Town & London. 
Macdonald, J. D. 1952. Notes on the Long-bill Lark, Certhilauda curvirostris. Ibis, 1952: 

122-127. 
Meinertzhagen, R. 1951. Review of the Alaudidae. Proc. zool. Soc. Lond. 121: 81-132, 

6 text-figs. 
Roberts, A. 1936a. Ornithological Notes. Ann. Transv. Mus. 18: 255-269. 
19366. Some unpublished Field Notes made by Dr. (Sir) Andrew Smith. Ann. Transv. 

Mus. 18: 271-323. 
1937- Some results of the Barlow-Transvaal Museum Expedition to South -West Africa. 

Ostrich, 8: 84-111. 

1940. Birds of South Africa. . . . xxxii + 463 pp., 56 pis. col. London & Johannesburg. 

Sclater, W. L. 1911-1912. On the birds collected by Mr. Claude H. B. Grant in various 

localities in South Africa. Ibis, 1911: 208-437, 695-741 ; 1912: 1-63. 

1930. Systema Avium Aethiopicarum. Pt. II. 305-922. London (Brit. Orn. Union). 

Sharpe, R. B. 1874. A study of the larks of southern Africa. Proc. zool. Soc. Lond. 1874: 

614-651. 
Smith, A. 1843. Illustrations of the Zoology of South Africa. ... 2, London. 
Strickland, H. E. 1852. List of a collection of birds procured by C. T. Andersson in the 

Damara country of South-western Africa. Jardine's Contribution to Ornithology 1852: 

141-160. 
Villalobos, C. & J. 1947. Colour Atlas. Buenos Aires. 
Vincent, A. W. 1946. On the Breeding Habits of some South African Birds. Ibis, 1946: 

462-477. 
Wallis, J. P. R. 1936. Fortune my Foe; the story of C. J. Andersson, African explorer, 18 37- 

1867. 312 pp. London. 



TABLES OF MEASUREMENTS 

Abbreviations: B.M. — British Museum 

T.M. — Transvaal Museum 
S.A.M. — South African Museum 
M.C. — Meinertzhagen Collection 
F.P. — First Primary 
V.C.— B.M. Vellum Catalogue 



C. a. albescens 



Locality 


Date 


Alt. 


Age 


Sex 


Wing 


Tail 


Bill 


F.P. 


Reference 


Cape of Good Hope* 


? 


? 


Ad. 


? 


90 


64 


17 


33 


B.M. 


>> 


? 


? 


Ad. 


? 


88 


63 


19 


3i 


B.M. 45:7:6:212 


>> 


? 


? 


Ad. 


? 


89 


63 


19 


27 


B.M. V.C. 18:19a. 


Cape Town 


? 


S.L. 


Ad. 


p 


85 


60 


18 


28 


B.M. 70:12:31:750 


Blaauwberg 


21:6 


S.L. 


Ad. 


6 


9i 


64 


19 


? 


B.M. 1937:7:14:238 


j> 


21:6 


S.L. 


Ad. 


$ 


88 


62 


19 


33 


B.M. 1937:7:14:237 


Milnerton 


? 


S.L. 


Ad. 


<J 


88 


64 


18 


32 


S.A.M. 13272 


> » 


? 


S.L. 


Ad. 


<J 


93 


67 


20 


32 


T.M. 20765 


>> 


p 


S.L. 


Ad. 


$ 


83 


56 


? 


30 


S.A.M. 13273 


„ 


? 


S.L. 


Ad. 


? 


82 


57 


18 


28 


S.A.M. 13273 


,, 


p 


S.L. 


Ad. 


$ 


84 


60 


19 


29 


S.A.M. 13273 


Durban Road 


11:4 


p 


Ad. 


3 


90 


62 


19 


28 


S.A.M. 14910 


Philadelphia 


10:4 


p 


Ad. 


6 


94 


68 


20 


30 


S.A.M. 


Berg River 


? 


? 


Ad. 


$ 


88 


61 


20 


27 


B.M. 76:5:23:705 


8 mis. NE. of Cape 




















Town 


26:4 


• 


Ad. 


? 


87 


60 


19 


32 


M.C. 



* Type of Alauda codea Smith. 



RED-BACK LARKS OF WESTERN SOUTH AFRICA 

C. a. saldanhae 



347 



Locality 


Date 


Alt. 


Age 


Sex 


Wing 


Tail 


Bill 


F.P. 


Reference 


Saldanha Bay* 


15:11 


S.L. 


Ad. 


8 


9i 


62 


19 


? 


T.M. 11881 


,, 


10:10 


S.L. 


Ad. 


8 


91 


66 


20 


32 


S.A.M. 7708 


,, 


10:10 


S.L. 


Ad. 


8 


9i 


68 


19 


31 


S.A.M. 7809 


Berg River 


p 


p 


Ad. 


? 


90 


64 


20 


29 


B.M. 76:5:23:706 


,, 


? 


p 


Ad. 


? 


93 


66 


20 


30 


B.M. 89:9:13:82 


Lambert's Bay 


29:10 


? 


Ad. 





92 


7i 


19 


? 


T.M. 11878 


Port Nolloth 


19:8 


S.L. 


Ad. 


8 


95 


7i 


20 


32 


B.M. 1950:52:12 


,, 


19:8 


S.L. 


Ad. 


8 


95 


72 


22 


? 


T.M. 20901 


t> 


19:8 


S.L. 


Ad. 


9 


87 


66 


18 


? 


T.M. 20899 


,, 


20:8 


S.L. 


Ad. 


$ 


85 


64 


19 


? 


T.M. 20902 


25 mis. E. of P. 




















NoUoth 


20:8 


500 


Ad. 


9 


86 


61 


20 


p 


T.M. 20903 



Swellendam 
Nels Poort 
Traka 
Deelfontein 



Berg-Orange Riverf 
Klaver 

van Rhynsdorp 



CalviniaJ 



40 mis. E. of C. 



Brandvlei 
Nr. Carnarvon 
Kamieskroon 



Grootberg 
Springbok 

m 
NW. of Springbok 



? 

p 

? 

23:2 

28:2 

28:1 

7:3 
p 

28:9 

28:9 

? 

? 

? 

?:8 

?:8 

30:4 

30:4 

30:4 

i:5 

1:5 

1:5 

30:4 

2:5 

2:5 

2:5 

2:5 

3:5 

12:6 

25:9 
28:9 
30:9 
2:12 
2:12 
4:12 
3:12 
3:12 
29:7 
29:7 
6:12 
7:12 
7:12 
7:12 
6:12 
8:5 
8:5 
8:5 



2,500 
2,500 
2,500 
2,500 
2,500 
2,500 
2,500 
2,500 



2,600 
2,600 
2,600 
2,600 
2,600 

? 

? 

? 



c. 


a. guttata 








Ad. 


? 


83 


62 


18 


30 


Ad 


? 


92 


64 


17 


26 


Ad. 


? 


9i 


68 


? 


34 


Ad. 


8 


94 


69 


? 


28 


Ad. 


? 


84 


62 


16 


26 


Juv. 


8 


92 


66 


15 


29 


Juv. 


9 


? 


? 


? 


? 


Ad. 


? 


90 


65 


19 


28 


Ad. 


8 


92 


66 


18 


? 


Ad. 


8 


93 


67 


18 


33 


Ad. 


8 


93 


65 


19 


30 


Ad. 


9 


85 


60 


18 


? 


Ad. 


9 


84 


60 


18 


29 


Ad. 


8 


99 


72 


19 


? 


Ad. 


9 


89 


64 


17 


? 


Ad. 


8 


93 


7i 


18 


35 


Ad. 


8 


95 


72 


18 


33 


Ad. 


8 


90 


67 


18 


33 


Ad. 


8 


96 


69 


? 


32 


Ad. 


8 


90 


66 


18 


32 


Ad. 


9 


85 


66 


18 


28 


Ad. 


? 


89 


65 


17 


26 


Ad. 


8 


90 


64 


18 


29 


Ad. 


8 


92 


66 


19 


29 


Ad. 


8 


94 


7i 


18 


3i 


Ad. 


9 


89 


66 


17 


30 


Ad. 


8 


94 


71 


18 


3i 


Ad. 


8 


90 


71 


17 


27 


Ad. 


8 


94 


68 


? 


32 


Ad. 


8 


96 


7i 


? 


32 


Ad. 


8 


92 


65 


19 


36 


Ad. 


8 


88 


63 


17 


32 


Ad. 


8 


93 


68 


18 


33 


Ad. 


8 


9i 


65 


19 


28 


Ad. 


9 


86 


60 


17 


27 


Juv. 


9 


86 


64 


18 


32 


Ad. 


8 


93 


67 


18 


36 


Ad. 


9 


84 


61 


? 


27 


Ad. 


8 


94 


68 


18 


34 


Ad. 


8 


100 


71 


19 


3i 


Juv. 


8 


89 


70 


17 


32 


Juv. 


8 


91 


74 


17 


28 


Ad. 


? 


89 


65 


19 


30 


Ad. 


8 


86 


7o 


19 


p 


Ad. 


8 


94 


7o 


19 


33 


Ad. 


8 


100 


74 


20 


3i 



B.M. 74:4:5:656 
B.M. 79:4:5:657 
79:4:5:658 
1903:3:9:470 
1903:3:9:471 
1901:9:5:23 
1901:9:5:26 
1845:7:6:213 
1 1 874 



B.M 
B.M 
B.M 
B.M 
B.M 
B.M 
T.M, 



S.A.M. 
T.M. 15211 



15210 
15209 
29012 
20913 



T.M. 

T.M. 

T.M. 

T.M. 

M.C. 

M.C. 

M.C. 

M.C. 

M.C. 

M.C. 

M.C. 

M.C. 

M.C. 

M.C. 

M.C. 

M.C. 

M.C. 

S.A.M. 18231 

S.A.M. 18233 

S.A.M. 18232 

B.M. 1950:50:940 

B.M. 1950:50:939 

B.M. 1950:50:934 

B.M. 1950:50:942 

B.M. 1950:50:941 

B.M. 73:10:20:145 

B.M. 73:10:20:232 

B.M. 1950:50:945 

B.M. 1950:50:946 

B.M. 1950:50:948 

B.M. 1950:50:947 

B.M. 1950:50:944 

M.C. 

M.C. 

M.C. 



* Type of Calendulauda 
t Type of Calendulauda 



albescens saldanhae 
guttata calviniensis 



Roberts. 
Roberts. 



■type of Alauda lagepa Smith. 



34« 



TAXONOMY OF THE KARROO AND 









C. a. guttata 


(cont.) 








Locality 


Date 


Alt. 


Age 


Sex 


Wing 


Tail 


Bill 


F.P. 


Reference 


NW. of Springbok 


8:5 


7 


Ad. 


a 


96 


72 


19 


36 


M.C. 


,, 


8:5 


7 


Ad. 


? 


90 


65 


18 


33 


M.C. 


,, 


8:5 


7 


Ad. 


? 


87 


63 


17 


30 


M.C. 


Klipfontein 


1:7 


3,000 


Ad. 


a 


88 


65 


18 


3i 


B.M. 1905:12:29:1437 


i> 


17:8 


3,000 


Ad. 


a 


9i 


68 


17 


7 


T.M. 20904 


>> 


22:12 


3,000 


Ad. 


$ 


92 


7 


18 


29 


B.M. 1950:50:952 


>> 


9:4 


3,000 


Ad. 


? 


85 


63 


17 


28 


B.M. 1905:12:29:1436 


>> 


5H 


3,000 


Ad. 


? 


90 


65 


i7 


28 


B.M. 1905:12:29:1435 


,, 


10:7 


3,000 


Ad. 


? 


87 


64 


18 


30 


B.M. 1905:12:29:1438 


,, 


22:12 


3,000 


Ad. 


? 


86 


60 


17 


30 


B.M. 1950:50:1000 


Anemous 


1:4 


600 


Ad. 


3 


90 


67 


7 


33 


B.M. 1905:12:29:1440 


>> 


1:4 


600 


Ad. 


a 


92 


63 


19 


28 


B.M. 1905:12:29:1439 


25 mis. E of P. 




















Nolloth 


19:8 


500 


Ad. 


3 


92 


67 


18 


7 


T.M. 20905 


>> 


19:8 


500 


Ad. 


a 


93 


66 


19 


7 


T.M. 20907 


,, 


19:8 


500 


Ad. 


a 


90 


62 


19 


7 


T.M. 20908 


„ 


19:8 


500 


Ad. 


a 


95 


66 


19 


7 


T.M. 20909 


>> 


22:8 


500 


Ad. 


a 


91 


62 


20 


7 


T.M. 2091 1 


,, 


21:8 


500 


Ad. 


? 


85 


61 


19 


7 


T.M. 20910 


>> 


19:12 


500 


Ad. 


? 


86 


60 


17 


28 


B.M. 1950:50:950 


f > 


19:12 


500 


Ad. 


? 


83 


59 


17 


29 


B.M. 1950:50:951 


>> 


19:12 


500 


Ad. 


? 


93 


66 


18 


29 


B.M. 1950:50:949 


Port Nolloth 


4:8 


S.L. 


Ad. 


a 


89 


63 


19 


30 


B.M. 1905:12:29:1442 


>> 


21:7 


S.L. 


Ad. 


? 


86 


58 


20 


30 


B.M. 1905:6:20:10 


»> 


18:7 


S.L. 


Ad. 


? 


88 


61 


18 


27 


B.M. 1904:6:20:9 



Grey Phase 

38 mis. N. of P. 

Nolloth 
Grootderm 



Orange Mouth 



Red Phase 
Grootderm 



Witputs 






C. a. patae 



19:12 
11:12 
13:12 
17:12 
17:12 
13:12 
17:12 
24:9 

?: 9 



13:12 
17:12 



23:1 

24:1 

24:1 

24:1 

24:1 

26:1 

25:1 

25:1 

23:1 

23 

25 

26 

26 

26 

27 

24 

25:1 

26:1 



Type 



S.L. 
500 
500 
500 
500 
500 
500 

S.L. 

S.L. 

S.L. 

500 
500 



4,000 
4,000 
4,000 
4,000 
4,000 
4,000 
4,000 
4,000 
4,000 
4,000 
4,000 
4,000 
4,000 
4,000 
4,000 
4,000 
4,000 
4,000 
of C. a, 



Ad. 
Ad. 
Ad. 
Ad. 
Ad. 
Ad. 
Ad. 
Ad. 
Ad. 
Ad. 

Ad. 
Ad. 



86 
89 
96 
92 
89 
84 
86 
86 
87 
82 

86 



C. a. cavei 



Ad. 
Ad. 
Ad. 
Ad. 
Ad. 
Ad. 
Juv. 
Juv. 
Ad. 
Ad. 
Ad. 
Ad. 
Ad. 
Ad. 
Ad. 
Juv. 
Juv. 
Juv. 
patae. 



93 
92 
95 
96 

93 
96 

89 
9i 
85 
83 
87 
85 
84 
86 

85 
78 
84 
82 



63 
? 

68 
64 
67 
61 
62 
64 
64 
62 

63 

67 



69 
69 
7o 
70 
7o 
7i 
65 
72 
64 
59 
64 
62 
64 
64 
63 
65 
64 
61 



20 
20 
20 
21 
21 
18 
17 
20 
19 
17 

21 
20 



27 
27 
26 
? 

26 
27 
30 
7 



29 
? 

30 

33 
7 



26 

19 
7 

28 
26 

7 

7 

29 
27 
30 

27 
28 



B.M. 
B.M. 
B.M. 
B.M. 
B.M. 
B.M. 
B.M. 
T.M. 
T.M. 
T.M. 



1950:50:938 

1950:50:937 

1950:50:933 

1950:50:934 

1950:50:936 

1950:50:932 

1950:50:935 

25274 

25275 

25276 



B.M. 1950:50:929 
B.M. 1950:50:930 



B.M. 
B.M. 
B.M. 
B.M. 
T.M. 
B.M. 
B.M. 
B.M. 
B.M. 
B.M. 
B.M. 
B.M. 
B.M. 
B.M. 
B.M. 
B.M. 
B.M. 
B.M. 



1950:50:928 

1950:50:926 

1950:50:913 

1950:50:924 

(C 205) 

1950:50:922 

1950:50:916 

1950:50:918 

1950:50:925 

1950:50:923 

1950:50:912 

1950:50:915 

1950:50:920 

1950:50:921 

1950:50:927 

1950:50:914 

1950:50:919 

1950:50:917 



t Type of C. a. cavei. 



RED-BACK LARKS OF WESTERN SOUTH AFRICA 

C. a. barlowi 



349 



Locality 


Date 


^/f. 


Age 


Sex 


Wing 


Tail 


Bill 


F.P. 


Reference 


Aus (scrub)* 


3i:7 


5 ,000 


Ad. 


<$ 


91 


68 


20 


? 


T.M. 20876 






3i:7 


5, 000 


Ad. 


c? 


95 


67 


20 


30 


B.M. 1950:52:10 






30:1 


5,000 


Ad. 


c? 


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B.M. 1950:50:908 






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B.M. 1950:50:901 






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B.M. 1950:50:911 






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30:1 


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B.M. 1950:50:909 






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5,000 


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86 


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1:2 


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89 


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C. a. erythrochlamys 



Kuiseb, nr. Walvis 








Bay 


6:9 


S.L.-600 


Ad. 




6:9 


S.L.-600 


Ad. 




18:11 


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? 


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18:12 


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20:11 


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Ad. 




26:5 


S.L.-600 


Juv 




? 


S.L.-600 


Juv 


Tsondab Mund 


5:3 


2,700 


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5:3 


2,700 


Ad. 




5:3 


2,700 


Ad. 




6:3 


2,700 


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6:3 • 


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24914 

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89:9:13:179 

73:18:20:218 

76:5:23:710 

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76:5:23:708 

66:7:19:2 

1950:50:898 

1950:50:882 

1950:50:886 

1950:50:893 

1950:50:890 

1950:50:891 

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1950:50:894 

1950:50:899 

1950:50:896 

1950:50:889 

1950:50:897 

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1950:50:885 

1950:50:895 

1950:50:892 

1950:50:887 

1950:50:884 

1950:50:881 

1950:50:877 

20888 

20885 

(R598) 

20893 

20889 



Type of Pseudammomanes barlowi Roberts. 



35Q 



TAXONOMY OF THE KARROO AND RED-BACK LARKS 



C. a. erythrochlamys (cont. 





Locality Date 


Alt. 


Age 


Sex 


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PRESENTED 

2 8 JAN 1953 



Bull. B.M. (N.H.) Zoology /, 11 



PLATE 36 



RACES AND DISTRIBUTION OF 

Certhilauda albescens 
IN WESTERN SOUTH AFRICA 




Bull. B.M. (N.H.) Zoology I, 11 



PLATE 37 







erythrochlamys 



barlowi 



patae 



guttata 



saldanhae 



albescens 



VARIATION IN CERTHILAUDA ALBESCENS 

For distribution of races see map. Changes in pattern are fairly evident and some changes 

in colour tone can be distinguished 



Bull. B.M. (N.H.) Zoology I, 11 



PLATE 38 




* A Hi 








■3^ . 




(a) Typical desert-edge country about five miles south of Witputs, Huns Mts., with sparse low 

scrub and prostrate succulents. Red-back Larks (Certhilanda albescens cavei) were present in 

twos ; spike-heel Larks [Certhilauda albofasciata) in parties of three to five ; and Red-cap Larks 

(Tephrocorys cinerea) in restless flocks 




(b) Shifting sand dunes in the Namib Desert at Tsondab Mund, showing clumps of spiky 
Aristida grass frequented by Red-back Larks (Certhilauda albescens erythrochlamys) 




PRESENTED 

2 8 JAN 1953 



PRINTED IN 

GREAT BRITAIN 

AT THE 

UNIVERSITY PRESS 

OXFORD 

BY 

CHARLES BATEY 

PRINTER 

TO THE 

UNIVERSITY 




3 DEC 1953 



SUBERITES DOMUNCULA (OLIVI): ITS 
SYNONYMY, DISTRIBUTION, AND ECOLOGY 

M. BURTON 



NOTES ON ASTEROIDS IN THE 
BRITISH MUSEUM (NATURAL HISTORY) 

III and IV 

A. M. CLARK 



SOME INTER-TIDAL MITES FROM 
SOUTH-WEST ENGLAND 

G. O. EVANS AND E. BROWNING 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ZOOLOGY Vol. i No. 12 

LONDON : 1953 



SUBERITES DOMUNCULA (OLIVI): ITS 
SYNONYMY, DISTRIBUTION, AND ECOLOGY 



BY 

M. BURTON 



NOTES ,ON ASTEROIDS IN THE 
BRITISH MUSEUM (NATURAL HISTORY) 

III and IV 



BY 

A. M. CLARK 



tf 



SOME INTER-TIDAL MITES FROM 
SOUTH-WEST ENGLAND 

BY 

G. O. EVANS AND E. BROWNING 



Pp. 351-422; Ph. 39-46; 30 Text-figures 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ZOOLOGY Vol. 1 No. 12 

LONDON : 1953 



THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted in 1949, is issued 
in five series corresponding to the Departments of the 
Museum, and an Historical Series. 

Parts appear at irregular intervals as they become 
ready. Volumes will contain about three or four hundred 
pages, and will not necessarily be completed within 
one calendar year. 

This paper is Vol. 1, No. 12, of the Zoology series. 



PRINTED BY ORDER OF THE TRUSTEES OF 
THE BRITISH MUSEUM 

Issued December 1953 Price One Pound 



SUBERITES DOMUNCULA (OLIVI): 
ITS SYNONYMY, DISTRIBUTION, AND 

ECOLOGY 

By MAURICE BURTON 
INTRODUCTION 

The names Suberites domuncula and Ficulina ficus have appeared almost consistently 
side by side in the literature for nearly 200 years. At times they have been treated as 
synonyms, and on such occasions Suberites domuncula has sometimes been given 
priority, at other times Ficulina ficus. Several major attempts have been made 
(Lendenfeld (1897), Topsent (1900), and Vosmaer (1933)) to put the histories, 
synonymies, and affinities of these two species in order. Each has failed for one 
reason or another. The present work is a comprehensive survey, made in the hope of 
achieving a reasonable stability. 

Since the earliest of the two names, Ficulina ficus, is here accepted as a synonym of 
Suberites domuncula, a restatement of its history is a first requisite. In the following 
pages are included, among other things, notes on individual references to the two 
species in question, as well as to their numerous synonyms, and this is followed by 
their arrangement in the usual synonymy list. The first of these tasks was, in fact, 
done by Vosmaer (1933) very completely, so that to all appearances its repetition is 
unnecessary. It is, therefore, essential to point out in what manner Vosmaer 's work 
has failed. To begin with, while he justifiably, as I think, regarded Suberites domun- 
cula and Ficulina ficus as synonyms, and included other names such as F. lutkenii 
within the scope of the species, he went too far. For example, he included Tethya 
prunum Costa, which is quite unrecognizable. He also included Suberites montiniger 
Carter, which belongs more properly to the genus Pseudosuberites, and Suberites con- 
cinnus Lambe, which is a Hymeniacidon. Secondly, he did not have the advantage of 
Topsent 's (1933) analysis of the early history of the specific name ficus. Thirdly, he 
included in his list every possible reference to any of these names, and many of them 
are so trivial that to include them in the synonymy list, already unwieldy, makes it 
completely overburdened. For example, if an author mentions one of these names 
merely in passing the name figures in his list of synonyms. I have checked carefully 
Vosmaer's pages and have eliminated all such trivial entries. Finally, he included 
certain other references without any justification. These are now given below. 

Alcyonium ficus , Hatchett, 1800: 355 : this is an account of certain simple chemical 
tests carried out on a marine organism. There is no information from which the 
organism could be identified, and nothing to suggest that it is a sponge. From the 
reactions obtained by the use of some at least of the chemical reagents it would 
appear to be an Ascidian (probably the Sea-fig as then understood, either Poly- 
clinum or Aplydium). 

Suberites compactus, Crivelli, 1863: 297 (sep. pag. 14), pi. vi, figs. 4-6: this is a 



354 SUBERITES DOMUNCULA (OLIVI) 

Suberites too inadequately described for the characters of the species to be deter- 
mined with accuracy. 

Halichondria virgultosa. Under this title Yosmaer, 1933: 441 lists Esper (1798), 
Lamarck (1813), Lamarck (1816), Lamouroux (1816), Blainville (1819), Lamouroux 
(1824), Lamarck (1836), Johnston (1842), Gray (1848), Duchassaing and Michelotti 
(1864). None of these authors is dealing with the species later described by Bower- 
bank as Hymeniacidon virgultosa and recognized by subsequent authors as a synonym 
of Suberites ficus. 

THE EARLY HISTORY OF FICULINA FICUS 
The early history of the sponge species, known today as Ficulina ficus and known 
for nearly 200 years under various generic (and often specific) names, is one of un- 
usual confusion. This arose largely from the fact that the 'sea-fig' of the Mediter- 
ranean is a sponge, and the 'sea-fig' in English usage is a Tunicate. The shape in 
both is very similar, and so long as the description of animal species depended on 
external appearances the mistake was bound to be perpetuated. An attempt was made 
to straighten out this early confusion by R. Hartmeyer (/. Mar. Biol. Ass. 10 (2): 
262-282, 1914). This comprehensive paper seems to have escaped the attention of 
those working upon the taxonomy of sponges, chiefly because it was not included in 
the Zoological Record under Section III (Porifera or Spongida). According to Hart- 
meyer's analysis (I.e. : 264), the species ficus was for the first time 'used in a binomial 
combination with the generic name Alcyonium, so that Pallas must be regarded as the 
author of that species, which must bear the name ficus'. Pallas takes the Alcyonium 
tuberosum forma ficus of Imperato (1599, Ital. p. 599, lat. p. 839), as the first repre- 
sentative of Ficulina ficus (Pallas) Autt., and presumably Imperato's description 
must serve as the type of the species. 

Topsent (1933 : 27), in analysing the history of F. ficus, takes a very different view ; 
but before proceeding to his main argument it is necessary to note what he says con- 
cerning Spongia ficiformis Poiret, which writers of the late eighteenth and early nine- 
teenth centuries have accepted as a synonym of what we have later known as Ficulina 
ficus. It will be convenient, therefore, to dispose of Spongia ficiformis. Here, in 
tabular form, are Topsent 's views: 

Alcyonium pulmonaria Ellis and Solander, 1786 = Ascidian. 

Spongia ficiformis, Poiret, 1789 = Petrosia ficiformis (Poiret). [Topsent points out 
that the sponge recorded by Guettard (1789, pi. iii), and which Poiret took to 
represent the same species, was rightly named Spongia usitatissima by Lamarck.] 

Spongia ficiformis, Gmelin, 1791 = Petrosia ficiformis (Poiret). 

Spongia ficiformis, Esper, 1794: 282 = Petrosia ficiformis (Poiret). 

Spongia bulbosum (partim), Esper, 1798, pi. xx, fig. 4 = Petrosia ficiformis (Poiret). 

Spongia ficiformis, Lamarck, 1816: 47 = Petrosia ficiformis (Poiret). 

Spongia ficiformis, Lamouroux, 1824 = Petrosia ficiformis (Poiret). 

Topsent (op. cit.) then goes on to remark: 'Aucune confusion n'etait permise entre 
Spongia ficiformis Poiret et les animaux qui furent appeles Alcyonium ficus. Ce que 
les auteurs anciens, comme Marsilli et Ellis, ont decrit, et dont Pallas et Linne ont 



SUBERITES DOMUNCULA (OLIVI) 355 

fait A.ficus, etait une Synascidia, la Pulmonelle figure de de Blainville, et Lamouroux 
l'a fort bien reconnu, en la rapportant aux genres Polyclinum Cuvier ou Aplydium 
Savigny. Mais il semble que l'Eponge lisse, grisatre a l'exterieur, spongieuse, jaune 
pale a l'interieur, avec oscule au sommet, qu'il prit pour la Spongia fwiformis Poiret, 
etait plutot une Ficulina, et ce qu'on appelle Ficulina ficus devrait peut-etre se 
nommer F.ficiformis (Lamouroux).' In other words, Topsent takes the view that all 
references to the so-called Ficulina ficus prior to Lamouroux (1824) are concerned 
with either Petrosia ficiformis Poiret or an ascidian. If this be so, then Alcyonium 
tuberosum forma ficus of Imperato must belong to one or the other, also. The only 
opinion opposed to this is the one expressed by Hartmeyer [I.e. : 264) that it is 'with- 
out doubt a sponge and has been identified by the spongologists as Ficulina ficus '. 
This has practically no value. It certainly is not 'without doubt a sponge'; and if 
' the spongologists ' have identified it as Ficulina ficus, they have merely done so by 
implication, by copying without question the earlier writers. And these Topsent has 
shown to be wrong in their identifications. 

It is proposed here to accept Topsent 's view, which best accords with my re- 
examination of the evidence. Moreover, as I hope to show, there is good reason to 
regard the so-called Ficulina ficus as a synonym of Suberites domuncula (Olivi). 
Since Olivi's publication antedates the use of Spongia ficiformis by fifty years, the 
ultimate name of the species can no longer be in doubt. 

This earlier confusion is, however, paralleled by the subsequent history of the 
species, though this time in a different sense. Ficulina ficus is obviously a close rela- 
tive of Suberites domuncula (Olivi) . Indeed, broadly speaking, the latter is a Ficulina 
ficus growing commensally with a hermit crab, and I have long held the opinion that 
the two species cannot be separated generically and may even be conspecific. It is 
in order to assess the value of this opinion that the following analysis is undertaken. 

CHRONOLOGICAL LIST OF REFERENCES TO FICULINA FICUS AND 

SUBERITES DOMUNCULA AND THEIR SYNONYMS, WITH BRIEF NOTES 

ON THEIR TAXONOMIC VALUE 

Alcyonium ficus, Pallas, 1766: 356: the species is established in a binominal com- 
bination, and is said to occur in the Mediterranean and on the English coast. [Top- 
sent (1933 : 27) accepts Pallas's specimen as an ascidian.] 

Alcyonium domuncula, Olivi, 1792: 241: the species is based on the figure 104 in 
Ginnani 1755. So far as a drawing of this kind can be relied upon, this would appear 
to be the well-known Suberites domuncula of subsequent authors. Presumably 
Ginnani's figure must be accepted as the holotype of the species. [The doubt implied 
here results from Topsent's (1900) diagnosis of Ficulina ficus. Under this name, as 
well as under Suberites domuncula, he includes specimens growing round mollusc 
shells inhabited by a Eupagurus. In other words, the sponge which everyone else has 
accepted as Suberites domuncula Topsent assigns partly to that species and, in com- 
pany with the free-growing forms, partly to Ficulina ficus. In doing this he gives a 
very restricted description of Suberites domuncula and recognizes its restricted dis- 
tribution (i.e. to the Mediterranean only). On the other hand, he does not make it 
precisely clear what differences he finds between the forms he recognizes as Ficulina 



356 SUBERITES DOMUNCULA (OLIVI) 

ficus growing on the Eupagurus-shell and Suberites domuncula. After studying his 
words closely it seems that his method of distinguishing between them rests on the 
characters of the ectosome, in addition to the absence of microscleres. In my opinion 
these are poor characters to be used in this connexion, but were their use to be 
upheld by subsequent investigation, then it would be impossible to say if Ginnani's 
figure represented Ficulina ficus or Suberites domuncula (sensu Topsent (1900) 
et seq.)] 

[Spongia suberosa, Esper, 1794: 266, pi. xli, figs. 1-2: has been accepted by some 
authors as a synonym of Suberites domuncula (Olivi), but it has nothing to do with 
either Alcyonium ficus or A. domuncula. Ehlers (1870) does not mention it, and 
although it has the habit of Halichondria bowerbanki, its identity is at present 
uncertain.] 

Alcyonium domuncula, Draparnaud, 1801: 169: notes on the living sponge, in 
which it is assumed that the specimens growing in association with a hermit crab 
[Suberites domuncula Autt.) belong to the same species as those growing on a Dromia 
(i.e. the Ficulina ficus Autt.). 

Alcyonium domuncula, Renier, 1804: xxv: nothing new. 

Alcyonium bidbosum, Esper, 1806: 41: typical examples of Olivi's species are 
figured and described, but without information on the internal structure. 

Alcyonium tuberosum, Esper, 1806, pi. xx: it seems that the author regarded this 
as a form of the preceding species. 

Alcyonium domuncula, Renier, 1807, pi. in: nothing new. 

Spongia domuncula, Bertoloni, 1810: 103: nothing new. 

Acyonium [sic] domuncula, Lamarck, 1815: 76: nothing new, except to reaffirm 
that the Mediterranean is the type-locality. 

Alcyonium compactum, Lamarck, 1815: 166: this is described by Topsent (1933: 
40) as Suberites domuncula (Olivi) (partim?). 

Alcyonium domuncula, Lamarck, 1816: 394: nothing new. 

Alcyonium compactum, Lamarck, 1816: 400: from the Atlantic, appears to be 
Suberites domuncula (Olivi). Spicules not mentioned. (See also Topsent, 1933: 40.) 

Spongia domuncula, Lamouroux, 1816: 38: nothing new. 

Alcyonium ficus, Lamouroux, 1816: 348: the author draws attention to the con- 
fusion between the sponge and the tunicate (see Hartmeyer, I.e.). Spicules not men- 
tioned. 

Alcyonium compactum, Lamouroux, 1816: 354: from the Atlantic. Spicules not 
mentioned. 

Spongia suberia, Montagu, 1818: 100: although the author gives an excellent 
description of the sponge, he does not say anything of its spicules. It is growing on 
univalve shells and is orange-yellow in life. It is clearly the animal generally accepted 
as Suberites domuncula (Olivi). 

Spongia domuncula, Bertoloni, 1819: 230: nothing new. 

Spongia suberia, Blainville, 1819: 130: nothing new. 

Lithumena domuncula, Renier, 1820, pi. iv: nothing new. 

Spongia suberosa, Gray, 1821 : 361 : merely gives a brief summary from Montagu 
(1818). 



SUBERITES DOMUNCULA (OLIVI) 357 

Alcyonium domuncula, Martens, 1824: 534: 'Auf dem Schlammgrund langs der 
westlichen Kuste haufig.' Found on hermit crabs and also on the carapace of Cancer 
dromia. 

Spongia domuncula, Lamouroux, 1824: 337: gives a summary of the literature to 
date, adding nothing new. 

Alcyonum [sic] ficus, Risso, 1826: 381: pear- or fig-shaped, up to 45 mm. long, 
grows in the ' Regions madreporiques ' and is an intense green in life. Spicules not 
mentioned. Possibly this is the ascidian. 

Alcyonum [sic] domuncula, Risso, 1826: 380: the author recognizes three varieties — 

Var. I. Rubro aurantio, flavo, coeruleo variegato. 

Var. II. Albo, poris oblongis, satis magnis et regulariter per superficiem sparsis. 

Var. III. Griseo et rubro aurantio variegato. Spicules not mentioned. 

I Halichondria suberica, Fleming, 1828: 522: mainly repeats Gray (1821) and adds, 
'I have found this species encrusting Corallines in the Firth of Forth.' The spicules 
are described as 'fusiform and slightly curved', the colour 'yellow'. 

Litumena spugnosa, Renier, 1828, pi. v ; nothing new. 

Anthelia domuncula, Blainville, 1830: 487: nothing new. 

Halichondria suberica, Coldstream, 1830 : 235 : two specimens from Rothesay Bay, 
on Turritella terebra. No colour notes and the only spicule figured is the tylostyle. 

Suberites domuncula, Nardo, 1833 : 523 ; nothing new. 

Suberites ficus, Nardo, 1833: 523: nothing new. 

Anthelia domuncula, Blainville, 1834: 524: nothing new. 

Suberites domuncula, Nardo, 1834: 714: nothing new. 

Spongia suberica, Lamarck, 1836 : 537 : nothing new. 

Alcyonium domuncula, Lamarck, 1836: 600: nothing new. 

Alcyonium compactum, Lamarck, 1836: 606: nothing new. 

Halispongia suberica, Blainville, 1837: 532: nothing new. 

Halichondria suberica and Spongia suberica, Thompson, 1840: 254: from Strangford 
and Belfast Loughs, 'investing univalve shells'. Spicules not mentioned. 

Halichondria suberica, Bellamy, 1840 : 268 : records the typical specimens, as well 
as those 'enveloping stems of sea-weed', from Devon. 

Halichondria suberea, Johnston, 1842: 139: adds little that is new. 

Halichondria ficus, Johnston, 1842 : 144 : deep water off Scarborough and Hartle- 
pool ; pear-shaped or rounded, often growing on shells ; greyish-white ; no mention of 
microscleres. 

Halichondria domuncula, Gray, 1848: 13: nothing new. 

Halichondria ficus, Gray, 1848: 15: nothing new. 

Halichondria suberea, Bowerbank, 1858: 287: gives the first good drawing of the 
megasclere. 

Halichondria ficus, Bowerbank, 1858: 298: the strongylote microsclere is figured. 

Halichondria compacta, Lieberkuhn, 1859: 520: on Buccinum and Murex inhabited, 
usually, by Pagurus callidus ; colour of red-lead ; spicules tylostyli. 

Halina suberea, McAndrew, 1861 : 235 : nothing new. 

1 This seems to contain the first mention of spicules, but megascleres only are mentioned. The first 
mention of microscleres is in Bowerbank, 1858. 



358 SUBERITES DOMUNCULA (OLIVI) 

Halina ficus, McAndrew, 1861: 235: nothing new. 

Hymeniacidon subereum, Bowerbank, 1862: 1111: nothing new. 

Halichondria ficus, Bowerbank, 1862: 1129: 'An elongated form of Halichondria 
ficus has also been again described as H. virgultosa' (i.e. by Johnston, 1842). 

Suberites domuncula, Schmidt, 1862: 67: largely reiterates Lieberkiihn's notes, but 
adds that there are two varieties, one from Quarnero which ' hat vorwiegend stumpf e 
Nadeln' (? = microstrongyla) , and the other, from Zlarin which 'hatte eine ganz 
prachtige Farbung, indem sie auf weissem und rothem Grunde lazurblau gezeichnet 
war'. Schmidt also described the species as common and well known. 

Suberites domuncula, Crivelli, 1863 : 286 : notes and coloured pictures. 

Suberites domuncula, Kolliker, 1864: 71: nothing new. 

Halichondria ficus, Bowerbank, 1864: 222 [also as Hymeniacidon ficus p. 244]: the 
centrotylote microstrongylote is figured, otherwise nothing new. 

Hymeniacidon suberea, Bowerbank, 1864: 231: nothing new. 

Halichondria {Hymeniacidon) suberea, Hughes, 1866: 86: notes on the development 
of the gemmules. 

Hymeniacidon virgultosa, Bowerbank, 1866: 193: a number of specimens from the 
Dogger Bank, erect (?), subcylindrical and substipitate, the base enclosing a Fucus, 
Zoophyte, or Dentalium, and ranging from 2f in. to 15 in. in length and up to J in. 
diameter. The colour, dried, is light buff-yellow. 

Bowerbank's specimens do not belong to the same species as ^-Halichondria vir- 
gultosa Johnston, which is apparently a Suberites sp. but of different habit ; nor, it 
may be presumed, to the Spongia virgultosa of Lamarck and Lamouroux. 

Hymeniacidon suberea, Bowerbank, 1866: 200: gives 'Locality. — The whole of the 
British coast', and 'colour. — Alive, yellow or orange; dried, yellow or brown'. His 
extensive notes show that he had difficulty in distinguishing between this species and 
Suberites carnosa on the basis of their respective spicules, and between Hymeniacidon 
suberea and Ficulina ficus on the basis of habitus. He found the species surrounding 
shells 'of Turbo, Fusus and other univalves', 'based on a Dentalium, a Vermetus, or 
some other equally ill-chosen locality', as 'large massive specimens', or 'partially 
enveloping a shell of a Fusus, the mollusc evidently alive at the time'. He also 
records ' a specimen as large as a hen's egg, attached by a broad base to the side of 
St. Katherine's Rock, at Tenby, between high and low water mark'. Bowerbank sees 
in the 'minute inflato-cylindrical ' spicules (i.e. microstrongyla) the chief means of 
distinguishing Ficulina ficus from Hymeniacidon suberea. 

Hymeniacidon ficus, Bowerbank, 1866: 206: specimens from Scotland, Northumber- 
land, and Hebrides, coloured grey, white, or russet red when alive. The specimens 
ranged from encrusting on a Pecten shell, covering ' a small univalve shell precisely 
after the manner of H. suberea' , to bulbous or fig-shaped. Clearly Bowerbank 
has used the presence of microstongyla as a distinctive character, but finds some 
difficulty in distinguishing between H. ficus and H. suberea on the grounds of 
habitus. 

Halichondria farinaria, Bowerbank, 1866: 269: is encrusting on Pecten opercularis, 
from Belfast Bay, Firth of Clyde and off Hastings, at 5 fathoms. It is scarlet or 

1 See last paragraph of the introduction (above). 



SUBERITES DOMUNCULA (OLIVI) 359 

reddish-orange in life and seems to have been found in fair numbers in the dredges. 
Microstrongyla are present. 

Renieraficus, Schmidt, 1866 : 16 : it is (erroneously) suggested that this is a synonym 
of R. (Hymeniacidon) caruncula. 

Suberites /urinaria, Schmidt, 1866: 16: nothing new. 

Reniera virgultosa, Gray, 1867: 518: nothing new. 

Halichondria f armaria, Gray, 1867: 519: nothing new. 

Suberites suberea, Gray, 1867 : 523 : nothing new. 

Ficulina ficus, Gray, 1867 : 523 : nothing new. 

Suberites domuncula, Marcusen, 1867: 358: from the Black Sea. 

Hymeniacidon subereus, Norman, 1868: 331 : from the Shetlands. 'Not so common 
as M. [sic] ficus, to which it is very closely allied.' 

Hymeniacidon ficus, Norman, 1868: 331: from the Shetlands. 'Common, coating 
univalve shells, and generally inhabited by hermit crabs/ 

Suberites suberia, Parfitt, 1868: 12: common along the Devon coast. No other 
information. 

Suberites domuncula, Schmidt, 1868: 14: gives a faunistic record for Algeria, with- 
out other comment. 

Halichondria farinaria, Bowerbank, 1868: 124: nothing new. 

Hymeniacidon suberea, Wright, 1869 : 53 : nothing new. 

Halichondria farinaria, Wright, 1869: 54: nothing new. 

Hymeniacidon ficus , Norman, 1869: 297: from Oban. 

Halichondria suberea, Carter, 1870 : 82 : notes on the gemmules. Carter considers 
the sponge has the property of dissolving shells and places it in the Clioniadae (of 
Gray) . 

Suberites heros, Schmidt, 1870 : 46 : a sponge from the Antilles, with the habitus of 
5. domuncula, 'ij Faust gross', and spicules ranging from styli to sub ty lost yli or 
tylostyli. 

Suberites lutkenii, Schmidt, 1870: 47: a new species, with microspined microscleres 
is described, from Denmark and Greenland. 

Suberites domuncula, Schmidt, 1870 : 76 : nothing new. 

Suberites ficus, Schmidt, 1870: 76: nothing new. 

Hymeniacidon virgultosa, Schmidt, 1870: 76: nothing new. 

Hymeniacidon suberea, Schmidt, 1870: 76: the author thinks this the same as 
Suberites domuncula. 

Halichondria farinaria, Schmidt, 1870 : yy : nothing new. 

Alcyonium domuncula, des Moulins, 1872 : 342 : the taking is recorded of this sponge 
in large numbers in fishermen's nets in the Gulf of Lyons. The hermit crab is extracted 
and used as bait. A synonymy list of the species is given. 

Suberites lutkenii, Mobius, 1873 : 148 : nothing new. 

Hymeniacidon ficus, Macintosh, 1874: 143: specimens, growing on Dentalium 
entalis, 'frequent on muddy ground'. 

Suberites lutkenii, Schmidt, 1874:429: nothing new. 

Hymeniacidon virgultosa, Bowerbank, 1874: 89: more specimens examined since 
1866, growing on univalve shells, and on a flat mass 'so like H. suberea that it is only 

zoo. 1, 12. 2 B 



360 SUBERITES DOMUNCULA (OLIVI) 

by microscopical examination that it can be separated from that species'. Micro- 
strongyla present. 

Hymeniacidon suberea, Bowerbank, 1874: 91: a specimen, from the Shetlands, in 
about 70 fathoms, of massive form enclosing a shell. 

Hymeniacidon ficus, Bowerbank, 1874: 92: more specimens, massive or ficiform, 
growing on bivalve shells or around univalve shells, from Tenby and the Island of 
Harris. Microstrongyla present. 

Halichondria f armaria, Bowerbank, 1874: 177: a small encrusting form, on Pecten 
opercularis, from Strangford Lough. Microstrongyla present. 

Suberites domuncula, Schmidt, 1875: 115: specimens from Solsvig, Peterhead, and 
Portobello, littoral to 50 fathoms. No other information. 

Suberites ficus, Schmidt, 1875: 116: a specimen from east of Bamborough, in 
36 fathoms on a bottom of sand and small stones. No other information. 

Halichondria suberea, Carter, 1875 : 197 : nothing new. 

Halichondria ficus, Carter, 1875: 197: nothing new. 

Suberites lutkenii, Liitken, 1875 : 190 : nothing new. 

Suberites domuncula, Carter, 1878 : 157 : nothing new. 

Suberites domuncula, Krukenberg, 1879: 66: notes on the physiology. 

Suberites domuncula, Krukenberg, 1879 : 705 : notes on the physiology. 

Suberites domuncula, Krukenberg, 1880: 37: notes on the physiology. 

Suberites montalbidus, Carter, 1880: 256: preliminary notice of a sponge from 
Barents Sea having centrotylote microxea for microscleres. 

Suberites domuncula, Czerniawsky, 1880 : 236 : from the Black Sea. 

Suberites domuncula, Leslie and Herdman, 1881: 60: nothing new. 

Halichondria suberea, Carter, 1881 : 255 : nothing new. 

Suberites domuncula, Vosmaer, 1881: 4: nothing new. 

Hymeniacidon virgultosus, Bowerbank, 1882 : 83 : nothing new. 

Hymeniacidon subereus, Bowerbank, 1882 : 88 : nothing new. 

Hymeniacidon ficus, Bowerbank, 1882: 89: abundant in Shetlands, Durham 
(Coralline zone), and specimens also from Oban ('on a pebble between tide-marks') 
and Westport, Co. Mayo. 

Halichondria farinaria, Bowerbank, 1882: 114: nothing new. 

Suberites domuncula, Klebs, 1882 : 295 : ' Der Schwamm . . . lebt stets auf Schnecken- 
schalen, in denen ein Pagarus lebt ; er umwachst die Mundung der Schale, so dass der 
Krebs haufig ganz eingeschlossen wird und sterben muss.' 

Halichondria suberia, Carter, 1882: 353: nothing new. 

Halichondria ficus, Carter, 1882 : 353 : nothing new. 

Suberites montalbidus, Carter, 1882: 353: a specimen from Barents Sea, with 
microstrongyla and faintly spined microxea, both centrolylote. 

Suberites domuncula, Graeffe, 1882: 318: from Trieste, with notes on ecology. 

Suberites domuncula, Vosmaer, 1882 : 20 : nothing new. 

Suberites sp., Vosmaer, 1882: 32: a specimen from the Arctic approximating to 
S. montalbidus. 

Suberites domuncula, Carter, 1883: 30: 150 specimens dredged 20 miles off 
Budleigh Salterton, growing on Turritella and Buccinum, with Pagurus or an 



SUBERITES DOMUNCULA (OLIVI) 361 

annelid inside, had incorporated much debris from the sea-bed in their sub- 
stance. 

Suberites domuncula, Marion, 1883 : 65 : notes, especially on its abundance, of the 
sponge off the Marseilles coast. 

Suberites domuncula, Vosmaer, 1884: 121: nothing new. 

Suberites domuncula, Vosmaer, 1885 : 332 : nothing new. 

Suberites montalbidus, Fristedt, 1885 : 19: records from the Swedish coast, in 75 m., 
of sponges with the spiculation shown by Carter (1882). 

Suberites ficus, Fristedt, 1885: 20: specimens from coast of Sweden, pale red in 
life, from various depths. Micros trongyla present. 

Suberites virgultosa, Fristedt, 1885: 21: five specimens from the Swedish coast, 
from unknown depths. Micros trongyla present. 

Suberites suberia, Higgin, 1886: 86: nothing new. 

Suberites domuncula, Vosmaer, 1886: 86: nothing new. 

Suberites domuncula, Vosmaer, 1886: 457: nothing new. 

Suberites liitkenii, Marenzeller, 1886: 3: the species is regarded as identical with 
S. montalbidus. 

Suberites montalbidus, Fristedt, 1887 : 428 : a number of specimens from Bering Sea 
and Bering Strait, the Siberian Arctic Ocean, Beaufort's Sea, Kara Sea, Barents Sea, 
and west of Greenland, in 2 to 40 fathoms, all having centrotylote microstrongyla and 
faintly spined microxea. 

Suberites domuncula, Ridley and Dendy, 1887: xlv: notes on histology. 

Suberites domuncula, Sollas, 1888 : 415 : notes on the structure of the skeleton. 

Suberites compactum, Topsent, 1888: 134: the sponge recorded by Lamouroux is 
said to be the equivalent of ' Spongia domuncula [Suberites ficus) \ 

Suberites domuncula, Topsent, 1888 : 134 : nothing new. 

Suberites ficus, Topsent, 1888: 134: is said to have the same Amphipod symbiont as 
5. domuncula. 

Suberites suberea, Topsent, 1888 : 150 : dredged at Luc and le Quihoc, it is encrust- 
ing and a deep orange. 

Suberites ficus, Topsent, 1888: 150: not common at Luc, it has the same habitat as 
5. suberea, and though orange-red as a rule, it is subject to 'decolorations partielles' 
and is often yellow or greyish. The surface is often perforated where an Amphipod, 
Tritacta gibbosa, is living. 

Suberites domuncula, Lendenfeld, 1888: 65: similar in habitat to the European 
forms, but although enclosing a crab the Australian forms do not contain shell with 
Pagurus. Colour bright yellow. Without microstrongyla. 

Suberites domuncula, Dendy 1889: 23: nothing new. 

Suberites domuncula, Lendenfeld, 1889 : 798 : is usually carried on the carapace of 
a Dromia. 

Suberites suberea, Hanitsch, 1889 : 158 : from Liverpool district. 

Halichondria farinaria, Topsent, 1889: xxxviii: nothing new. 

Suberites domuncula, Topsent, 1890 : 232 : nothing new. 

Suberites domuncula, Topsent, 1890: 232: 'partout dans la Manche/ 

Suberites suberea, Topsent, 1890 : 202 : from Luc. 



362 SUBERITES DOMUNCULA (OLIVI) 

Suberites ficus , Topsent, 1890: 202: from Luc. 

Suberites /armaria, Topsent, 1890 : 203 : nothing new. 

Suberites domuncula, Hanitsch, 1890: pp. 195, 214: gives records for the estuary of 
the Mersey, north Wales, Isle of Man, and Puffin Island, and declares that it may be 
found growing on bivalve shells and other substrata, as well as on univalve shells 
inhabited by hermit crabs. 

Suberites ficus, Hanitsch, 1890: 195: from north Wales. 

Suberites domuncula, Hanitsch, 1891: 218: several specimens from 10 fathoms off 
the west coast of Ireland. Hanitsch draws attention to the presence of microstrongyla, 
and to so many previous authors having missed them. 

Suberites ficus, Hanitsch, 1891: 219: two specimens from off the west coast of 
Ireland, in 5 to 15 fathoms. 

Suberites ficus, Topsent, 1891: 529: dredged at Roscoff. 

Suberites ficus, Topsent, 1891: 127: from Arcachon. 

Suberites ficus, Topsent, 1891: 14: two specimens from between Dakar and 
Rufisque, at 25 m., on muddy sand, with microstrongyla that lack a centrum. 

Suberites domuncula, Topsent, 1891 : 15 : a single littoral specimen from Dakar. 

Suberites domuncula, Topsent, 1891 : 15 : from Dakar. 

Suberites ficus, Topsent, 1891: 127, 129: from Arcachon. 

Suberites ficus, Topsent, 1891: 529: from Roscoff. 

Suberites domuncula, Hanitsch, 1891 : 218 : several specimens from the west coast in 
10 fathoms. He mentions the presence of centrotylote microstrongyla. 

Suberites ficus, Hanitsch, 1891: 219: from the west coast of Ireland in 5 to 15 
fathoms. 

Suberites latus, Lambe, 1892: 71: four specimens from British Columbia, lobo- 
massive, up to 60 mm. across, yellowish-brown in spirit, but without microstrongyla. 
Lambe (1893: 126) agrees this is conspecific with 5. suberea {= ficus). 

Suberites domuncula, Holt, 1892: 239: from Blacksod Bay, in 7 fathoms, on fine 
sand. 

Suberites ficus, Topsent, 1892: 128: four specimens from the Bay of Biscay in 
depths varying from 63 to 180 m. No mention is made of colour or the presence of 
microstrongyla. 

Suberites ficus, Levinsen, 1893: 410: numerous specimens from the Kattegat. 
According to the figures given, the spiculation resembles closely that of 5. montal- 
bidus. 

Suberites farinarius, Levinsen, 1893 : 412 : a specimen from the Kattegat, with 
centrotylote microscleres. 

Suberites montalbidus, Levinsen, 1893 : 413 : three specimens from the Kattegat in 
17 \ fathoms, showing the spiculation described by Carter (1882). 

Suberites domuncula, Celesia, 1893 : 1 : extensive notes on the relation between the 
form of the sponge and the presence of the hermit crab. 

Suberites ficus , Topsent, 1894: 21: from the Pas-de-Calais. Halichondria farinaria 
and H. virgultosa are regarded as synonyms. 

Suberites domuncula, Topsent, 1894: 23: from the Pas-de-Calais. 

Suberites suberea, Lambe, 1894: 126: nearly sixty specimens from Alaska. '. . . the 



SUBERITES DOMUNCULA (OLIVI) 363 

flesh-spicules are present in the majority of cases, but absent in a few; in some 
specimens they occur in great abundance, in others only one or two were seen. Evi- 
dently the presence or absence of the flesh-spicules cannot be considered of specific 
value.' 

Suberites montalbidus, Lambe, 1894: 127: a single example, 25 mm. across, from the 
Aleutians, with microscleres as described by Carter (1882). 

Suberites ficus, Weltner, 1894: 327: four specimens from the North Sea, including 
the Dogger Bank, from depths varying from 32 to 50 m. No colour records are given 
and microstrongyla are not mentioned. 

Suberites virgultosa, Hanitsch, 1894: 177: nothing new. 

Suberites domuncula, Hanitsch, 1894: 177: nothing new. 

Suberites ficus , Hanitsch, 1894: 177: nothing new. 

Suberites farinarius, Hanitsch, 1894: 179: nothing new. 

Suberites heros, Weltner, 1894: 328: suggests the identity of this species with 
5. ficus. 

Suberites suberea, Lambe, 1895: 126: records 60 specimens from Alaska, and points 
out (p. 127) that his 5. lotus, from Vancouver Island, is identical with 5. suberea. 

Suberites montalbidus, Lambe, 1895: 127: from Alaska. 

Suberites domuncula, Heider, 1895 : 283 : nothing new. 

Suberites ficus, Lambe, 1896: 193: two dried specimens from Nova Scotia, with 
microstrongyla, the one growing on a Pecten tenuicostata shell, the other on the inside 
of a shell of Cyprina. 

Suberites ficus, Topsent, 1896: 275: several specimens from the Bay of Biscay at 
140 to 400 m. 

Suberites ficus, Topsent, 1896: 118: from Quiberon (Atlantic coast of France). 

Ficulina ficus, Lendenfeld, 1896: 94: an extensive review of previous knowledge, 
with little additional information. 

Suberites domuncula, Lendenfeld, 1896: 118: a review of previous knowledge, with 
little additional information. 

Ficulina ficus, Topsent, 1898 : 129 : nothing new. 

Suberites heros, Thiele, 1898: 37: is probably identical with 5. domuncula. 

Suberites domuncula, Thiele, 1898: 37: the author differentiates between S. domun- 
cula, without microstrongyla, and 5. subereus, with microstrongyla (but see Lambe, 
1894: 126). 

Suberites liltkenii, Thiele, 1898: 38: is probably identical with 5. domuncula. 

Suberites subereus, Thiele, 1898 : 38 : several specimens from Japan, some enclosing 
shells, examined dry. Microstrongyla present. 

Suberites placenta, Thiele, 1898: 39: a depressed cake-shaped sponge from Japan, 
dry, with tylostyli and microstrongyla. 

Suberites sericeus, Thiele, 1898 : 39 : dry incrustations from Japan on a Pecten and 
a gastropod- shell, without microstrongyla, probably represent either 5. ficus or 
5. domuncula. 

Prosuberites inconspicuus , Thiele, 1898 : 40 : a dry encrusting specimen from Japan, 
in 100 fathoms, with tylostyli as in Thiele's specimen of Suberites subereus, but 
without microstrongyla, is probably a young 5. domuncula. 



364 SUBERITES DOMUNCULA (OLIVI) 

Prosuberites exiguus, Thiele, 1898: 40: two dried encrusting specimens from Japan, 
very like P. inconspicuus , probably represent young forms of Suberites domuncula. 
They are without microstrongyla. 

Ficulina ficus, Topsent, 1899 : 105 : recorded for the coast of Belgium without 
further details. 

Ficulina ficus, Topsent, 1900: 203: in a review of the species the author increases 
the confusion by using the presence or absence of the microstrongyla as a basis for 
the specific distinction. Consequently, under F. ficus are included all forms having 
microscleres regardless of the external form. 

Suberites lutkenii, Topsent, 1900: 213: is regarded as a variety of Ficulina ficus. 

Suberites domuncula, Topsent, 1900: 225: the species is interpreted in a narrow 
sense, depending almost entirely on the absence of microscleres. 

Suberites suberea, Lambe, 1900 : 161 : nothing new. 

Suberites ficus, Lambe, 1900: 161: nothing new. 

Suberites montalbidus , Lambe, 1900 : 162 : nothing new. 

Suberites montalbidus, Lambe, 1900: 24: from Hudson Bay and Strait. 

Suberites montalbidus, Lambe, 1900 : 277 : nothing new. 

Suberites domuncula, Cotte, 1901 : 1 : chemico-physiological notes. 

Suberites domuncula, Cotte, 1901 : 95 : physiological notes. 

Suberites domuncula, Bidder, 1902 : 380 : the author suggests that texture is a result 
of ecological conditions. 

Ficulina ficus, Rousseau, 1902 : 18 : the author treats Suberites domuncula as a 
synonym of this species and records it from the coast of Belgium. 

Suberites heros, Thiele, 1905 : 415 : nothing new. 

Suberites domuncula, Thiele, 1905 : 416 : nothing new. 

Suberites domuncula, Swartschewsky, 1905: 35: the species is recorded from the 
Black Sea. 

Suberites heros, Swartschewsky, 1905 : 35 : is accepted as a synonym of 5. domuncula. 

Suberites montalbidus, Swartschewsky, 1906: 318: from the White Sea. 

Ficulina ficus, Lundbeck, 1907: 558: 'Trois petits exemplaires pedunculeV. No 
other information. 

Ficulina ficus, Lundbeck, 1909 : 453 : one specimen, 100 mm. across, from East 
Greenland, in 25-40 fathoms. No other details. 

Ficulina ficus, Stephens, 1912: 21: the author accepts the identity of Suberites 
domuncula with this species and gives records for south-west Ireland from between 
tide-marks down to 8 fathoms. Massive specimens were found in littoral zone, and 
dredged specimens were growing on Pecten or on gastropod shells containing Eupa- 
gurus cuanensis. 

Ficulina ficus, Topsent, 1913 : 25 : from Norway ; a score of specimens 'enveloppant 
des coquilles et abritant des Pagures'. 

Ficulina lutkenii, Topsent, 1913: 25: from Norway. 

Ficulina ficus, Miiller, 1913: 291: the author treats Suberites domuncula and 
Ficulina ficus as one and the same thing. He gives notes on the gemmules in 
373 specimens from the Barents Sea, taken in 60-67 m - m August. Of this total 
261 were on bivalve shells, 6 on gastropod shells, and 36 on stones. The rest were 



SUBERITES DOMUNCULA (OLIVI) 365 

without point of attachment. Colour notes are not given, but microstrongyla are 
figured. 

Ficulina ficus, Stephens, 1915: 35: the author lists many records from Ireland. 

Suberites domuncula, Babic, 1921: 14: merely records the species for the Adriatic. 

Suberites domuncula, Babic, 1922: 272: several specimens, on Turritella, from the 
Adriatic, the largest 90 mm. in diameter. No colour records are given and no men- 
tion made of microstrongyla. 

Ficulina ficus, Ferrer, 1922: 269: nothing new. 

Suberites domuncula, Topsent, 1925 : 633 : records the species as common at Naples 
and varied in colour. He gives the opinion that the specimens at Naples do not attain 
such large proportions as those at Banyuls. 

Suberites domuncula, Dembowska, 1926: 163: an account of the habits of Dromia 
vulgaris and its use of the sponge. 

Ficulina ficus, Broch, 1927: 5: from Norway, Lindesness, in 20-24 m., growing 
on black mud. No other information. 

Ficulina ficus, Topsent, 1928: 156: specimens recorded from the Bay of Biscay and 
the Azores, from depths of 130 to 1,331 m. No colours are mentioned, and as to ex- 
ternal form the author merely says, of the specimens from Stn. 3660, that they are 
enveloping the shells of Gastropods. As to the specimen from a depth of 1,331 m., the 
author speaks of it as ' bien typique, amicrostrongyles centrotylotes, lisses, abondants \ 

Suberites domuncula, Topsent, 1928: 154: the species is recorded from off Toulon, 
in 20 m., with no other comment. 

Ficulina ficus, Arndt, 1928 : 33 : treats this species and Suberites domuncula as 
synonyms, and summarizes the characters of the species. 

Ficulina ficus, Hentschel, 1929: 928: nothing new. 

Ficulina lutkenii, Hentschel, 1929 : 928 : nothing new. 

Suberites domuncula, Burton, 1932: 201: a single specimen from Japan, in 10 
fathoms, enclosing a hermit crab. The synonymy of this species and Ficulina ficus is 
suggested. 

Suberites domuncula, Vosmaer, 1933 : 426 : a very extensive review of the species, 
but more confusion is caused by ascribing too wide limits to the species. 

Suberites domuncula, and Ficulina ficus, Burton, 1934: 313: the two species are 
compared. 

Ficulina lutkenii, Burton, 1934: 14: from East Greenland, at 3-191 m. 

Suberites domuncula, Topsent, 1934: 14: from Monaco. 

Ficulina ficus, Topsent, 1934: 16: in his specimens from Monaco, Topsent finds the 
occurrence of microstrongyla variable. In ' des cas embarrassants ' he succeeded ' par 
grattage du pourtour de l'oscule ' in finding a few in specimens which should otherwise 
be assigned to Suberites domuncula. 

Suberites domuncula, Arndt, 1935 : 39 : a summary of our knowledge of the species 
is given. 

Suberites ficus, Arndt, 1935: 39: in a summary of our knowledge of the species, 
Arndt returns to the orthodox method of distinguishing between this species and 
5. domuncula (i.e. basing his distinction solely on the presence or absence of micro- 
strongyla). 



366 SUBERITES DOMUNCULA (OLIVI) 

Suberites domuncula, Burton, 1935: JJ'. from the Sea of Japan, in 10-35 m - 
Suberites domunculus, de Laubenfels, 1949: 20: from Wood's Hole. The author 
appears to accept the identity of Ficulina ficus with Suberites domuncula. 

It would seem unnecessary to go into such minute detail, but for the confusion 
which has arisen independently of that caused by the early authors. In the main, 
authors since Lamouroux have treated as Ficulina ficus those specimens, with tylo- 
styli and centrotylote microstrongyla, growing with their bases implanted on a shell 
or other substratum. They have treated as Suberites domuncula any specimen of 
comparable structure completely enclosing a gastropod shell containing a hermit 
crab. Yet both species have the same two categories of spicules arranged in the same 
way, have a similar texture and colour, and have a similar geographical range and 
bathymetric distribution. These things have been recognized by Martens, Stephens, 
Arndt, and Miiller, who have regarded the two forms as conspecific. Admittedly 
these four authors form a minority, but it is worth recalling that Miiller examined 
373 specimens in a single investigation, and Stephens, whose work is of a uniformly 
high standard, must have handled more than this number in the course of a few years. 
I am the more inclined to accept their verdict since it coincides with my own (1934) 
arrived at independently. Against this we must set the views of many authors of 
limited experience, as well as those of Lendenfeld and, more especially Topsent, both 
workers of wide experience. Moreover, Arndt (1935) subsequently reverted to this 
view, apparently. The value of Lendenfeld's opinion can, however, be judged from 
his most extensive work on these two supposed species. In 1897 he set forth their 
characters in great detail and his figures show in each case that he was dealing with 
specimens enclosing a gastropod shell containing a hermit crab. In other words, he 
clearly had accepted the presence or absence of the microstrongyla as of specific 
importance. In Topsent 's (1900) main study of the two supposed species it is evident 
that he has adopted a similar plan. Lendenfeld, at least, seems to have based his 
action on Bowerbank (1866), who, while admitting the difficulty of distinguishing 
between the Ficulina ficus and Suberites domuncula, adopted the presence or absence 
of microstrongyla for their separation. It will be possible to show, not only that the 
presence or absence of the microstrongyla has no taxonomic value, but that at the 
most these two supposed species are probably no more than ecological varieties, if 
indeed there is that much separation. 

The history of the microstrongyla is quite remarkable. Although Suberites domun- 
cula was first described in 1792, it was not until 1828 that any mention of its spicules 
is made. Then Fleming described them as ' fusiform and slightly curved \ It was not, 
however, until 1834 that Coldstream figured a recognizable tylostyle. These are, 
however, the megascleres. No mention was made of the microscleres until much 
later, when Bowerbank (1858, p. 298) mentioned the finding of an 'inflato-cylindrical' 
in Halichondria ficus, and figured what is now called the centrotylote microstrongyle 
on pi. xxiv, fig. 25. In 1862 Schmidt wrote of 'stumpfe Nadeln', which may or may 
not refer to microstrongyla, and it was left to Lambe (1894), who examined nearly 
sixty specimens to show that they are present in Suberites domuncula as well as in the 
so-called Ficulina ficus. He found those microscleres present in varying numbers. In 



SUBERITES DOMUNCULA (OLIVI) 367 

only a few cases did he find them lacking in the typical Suberites domuncula. He pre- 
sumed, therefore, that ' the presence or absence of the flesh-spicules cannot be con- 
sidered of specific value \ Experience leads me to endorse Lambe's view ; and we may 
be reasonably sure that this is true also for workers such as Stephens and, possibly, 
Arndt. 

Another distinction that has been made between Suberites domuncula and Ficulina 
ficus is that the first is typically orange or red and the second typically green or 
greenish. Nobody has specifically stated this in print, but I have found it a prevalent 
opinion. If we summarize the colour records from the chronological list of references 
given above, we find that there is little to choose between them. Considering the 
number of times the two species have been referred to in the literature, colour records 
are meagre. They may be summarized as follows: 

Suberites domuncula : orange-yellow (Montagu) ; orange-red, white, grey and 
orange-red (Risso) ; yellow (Fleming) ; yellow or orange (Bowerbank) ; colour of 
red-lead (Lieberkuhn) ; white and red with blue patches (Schmidt) ; deep orange 
(Topsent) ; bright yellow (Lendenfeld) ; varied in colour (Topsent) ; usually 
orange, often white or white marbled with red and blue (Topsent) ; orange- 
yellow (Lendenfeld). 

Ficulina ficus : greyish- white (Johnston) ; scarlet or reddish-orange (Bowerbank) ; 
pale red (Fristedt) ; usually orange-red, often greyish or yellow (Topsent) ; 
orange-yellow (Lendenfeld) . 

It seems there is little to choose between the two forms in the matter of colour. 

The external form appears to have constituted a further barrier to recognizing the 
identity of Ficulina ficus with Suberites domuncula. In the former it is typically rig- 
or pear-shaped, with more or less of a stout peduncle, but variations are recognized 
up to the long, almost strap-shaped sponges seen in Bowerbank's Halichondria 
farinaria. The typical form in Suberites domuncula is oval or spherical with, on one 
side, an opening showing the presence of a hermit crab. What has not been recog- 
nized are the various intermediates between the two, and the fact that the association 
between the Suberites and the hermit crab is not a specific commensalism. To take 
the form first, Ficulina ficus has been recorded as growing on seaweeds and on bivalve 
and gastropod shells. It will, from my own observations, also grow on pebbles or rock 
surfaces. It may be encrusting, cushion-shaped, irregularly massive, lobose, ficiform, 
or elongated (f arinaria-f orm) . The base may surround to a varying extent the object 
to which it is attached. Suberites domuncula is normally encrusting, or spherical or 
subspherical, but may also be irregularly massive or lobose. The absence of the fici- 
form or elongated shape is almost certainly the result of the shell, on which the sponge 
is seated, being in a state of more or less continuous motion due to the presence in it 
of a hermit crab. 

That there is no specific commensalism between Suberites domuncula and a hermit 
crab may be shown by the following : 

The sponge has been found associated with : 

1. A wide variety of gastropod shells, which may often be without a hermit crab ; 

2. Several different species of Eupagurus ; 
zoo. 1, 12. 2 c 



368 SUBERITES DOMUNCULA (OLIVI) 

3. The carapace of a Dromia ; 

4. A Fusus, with the mollusc still alive. 

The evidence is markedly in favour of following the opinion of Arndt, Stephens, 
and others. There is, however, one point on which a reasonable doubt may be felt. 
This concerns the nature of Suberites montalbidus Carter. In the holotype its micro- 
scleres are microspined and centrotylote microxea in addition to the smooth centro- 
tylote microstrongyla. It seems, however, that this sharp distinction is not always 
maintained. Fristedt (1887), for example, also found both kinds in his Arctic speci- 
mens, but the microxea were but faintly spined and apparently not centrotylote. It 
is significant, nevertheless, that the recorded specimens of 5. montalbidus are from 
Barents Sea (Carter), Bering Sea and Strait, the Siberian Arctic, Kara Sea, Barents 
Sea, and west of Greenland (Fristedt) , Barents Sea (Levinsen) , and the Aleutians 
(Lambe), so that there is reasonable ground for suspecting that it constitutes a 
northern form. In the northern limits of its range Suberites domuncula {-\-Ficulina 
ficus) has also been recorded from Alaska, East Greenland, and Barents Sea. There 
is not, therefore, a clear line of geographical separation between it and S. montalbidus, 
and added to this Fristedt (1885) has recorded the latter from the coast of Sweden 
also. It may be that authors, such as Stephens, who have wide experience of S. 
domuncula, and have accepted 5. montalbidus as one of its synonyms, have found 
microspined microxea in southern individuals and have not considered it sufficiently 
important to draw attention to the fact. Under the circumstances, it would be better 
to follow the example set by experienced authors and regard 5. montalbidus as a 
synonym of 5. domuncula, at least for the present. 

REVISED LIST OF SYNONYMS OF SUBERITES DOMUNCULA, WITH A 
DESCRIPTION OF THE SPECIES, INCLUDING ITS DISTRIBUTION 

Suberites domuncula (Olivi) 

Alcyonium domuncula, Olivi, 1792: 241; Draparnaud, 1801: 169; Renier, 1804: 
xxv; A. bulbosum, Esper, 1806: 41; A. tuberosum, idem, I.e., pi. xx; A. domuncula, 
Renier, 1807: pi. iii; Spongia domuncula, Bertoloni, 1810: 103; Acyonium [sic] 
domuncula, Lamarck, 1815 : 76 ; Alcyonium compactum, idem, I.e. : 166 ; A. domuncula, 
idem, 1816: 394; A. compactum, idem, I.e. 400; Spongia domuncula, Lamouroux, 
1816: 38; Alcyonium ficus (partim?), idem, I.e. 348; A. compactum, idem, I.e. 
354; Spongia suberia, Montagu, 1818: 100; S. domuncula, Bertoloni, 1819: 230; 
5. suberia, Blainville, 1819: 130; 5. suberosa, Gray, 1821: 361; Alcyonium fici for me 
(partim?), Lamouroux, 1821: 29; A. domuncula, Martens, 1824: 534; Spongia 
domuncula, Lamouroux, 1824: 337; Alcyonium domuncula, Risso, 1826: 380; Hali- 
chondria suberica, Fleming, 1828: 522; Coldstream, 1830: 235; Anthelia domun- 
cula Blainville, 1830 : 487 ; Suberites ficus, Nardo, 1833 : 523 ; S. domuncula, idem, 
I.e.: 523; Anthelia domuncula, Blainville, 1834: 524; Halispongia suberica, idem, 
I.e.: 532; Suberites domuncula, Nardo, 1834: 714; Spongia suberica, Lamarck, 
I 836: 537; Alcyonium domuncula, idem, I.e.: 600; A. compactum, idem, I.e.: 606; 
Halichondria suberica, Bellamy, 1839: 268; Thompson, 1840: 254; H. suberea, 
Johnston, 1842: 139, pi. xii, figs. 5-6; H. ficus, idem, I.e.: 144, pi. xv, figs. 4-5; H. 



SUBERITES DOMUNCULA (OLIVI) 369 

domuncula, Gray, 1848 : 13 ; H.ficus, idem, I.e. : 15 ; H. suberea, Bowerbank, 1858 : 287, 
pi. xxiii, fig. 25 ; H. ficus, idem, I.e. : 298, pi. xxiv, fig. 25 ; H. compacta, Lieberkuhn, 
1859: 520; Halina suberea, McAndrew, 1861: 235; H. ficus, idem, I.e.: 235; 
Hymeniacidon subereum, Bowerbank, 1862: 1111; idem, I.e.: 1129; Suberites 
domuncula, Schmidt, 1862: 67; Crivelli, 1863: 286, pi. iii, figs. 1-5; Kolliker, 1864: 
71; Hymeniacidon ficus, Bowerbank, 1864: 222; H. suberea, idem, I.e.: 231, pi. i, 
fig. 23; H. virgultosa, idem, 1866: 193; H. suberea, idem, I.e.: 200; H. ficus, idem, 
I.e.: 206; H. farinaria, idem, I.e.: 269; Halichondria suberea, Hughes, 1866: 86; 
Reniera ficus, Schmidt, 1866: 16; Hymeniacidon farinaria, idem, I.e.: 16; Reniera 
virgultosa, Gray, 1867: 518; Halichondria farinaria, idem, I.e.: 519; Suberites 
suberea, idem, I.e.: 523; Ficulina ficus, idem, I.e.: 523; Suberites domuncula, 
Marcusen, 1867, p. 358; S. suberia, Parfitt, 1868: 12; Halichondria farinaria, Bower- 
bank, 1868: 124; Suberites domuncula, Schmidt, 1868: 14; Hymeniacidon ficus, 
Norman, 1869: 297 ; H. subereus, idem, I.e. : 331 ; H.ficus, idem, I.e. : 331 ; H. suberea, 
Wright, 1870: 225; Halichondria farinaria, idem, I.e.: 226; H. suberea, Carter, 
1870: 82; Suberites heros, Schmidt, 1870: 46; 5. lutkenii, idem, I.e.: 47, pi. v, fig. 7; 
5. domuncula, idem, I.e. : 76 ; 5. ficus, idem, I.e. : 76 ; Hymeniacidon virgultosa, idem, 
I.e.: 76; H. suberea, idem, I.e.: 76; Halichondria farinaria, idem, I.e.: 77; Alcyonium 
domuncida, Moulins, 1872: 342; Suberites lutkenii, Mobius, 1873: 148; Schmidt, 
1874: 429; Hymeniacidon virgultosa, Bowerbank, 1874: 89, pi. xxxv, figs. 1-5; H. 
suberea, idem, I.e. : 91, pi. xxxvi, figs. 1-4 ; H.ficus, idem, I.e. : 92, pi. xxxvi, figs. 10-17 i 
Halichondria farinaria, idem, I.e.: 177, pi. lxx, figs. 5-8; Hymeniacidon ficus, 
M'Intosh, 1874: 143; Halichondria suberea, Carter, 1875: 197; H. ficus, idem, I.e.: 
197; Suberites domuncula, Schmidt, 1875: 115; S. ficus, idem, I.e.: 116; 5. lutkenii, 
Lutken, 1875: 190; 5. domuncula, Carter, 1878: 157; Krukenberg, 1879: 66, pi. i, 
figs. 3-4 ; idem, 1879 : 705 ; Czerniawsky, 1880 : 236 ; Krukenberg, 1880 : 37 ; 5. montal- 
bidus, Carter, 1880 : 256 ; 5. domuncula, Leslie and Herdman, 1881 : 269 ; Vosmaer, 
1881 : 4 ; Halichondria suberea, Carter, 1881 : 255 ; Hymeniacidon virgultosa, Bower- 
bank, 1882: 83; H. subereus, idem, I.e.: 88; H.ficus, idem, I.e.: 89; Halichondria 
farinaria, idem, I.e.: 114; H. suberia, Carter, 1882: 353; H.ficus, idem, I.e.: 353; 
Suberites montalbidus, idem, I.e. : 353 ; 5. domuncida, Graeffe, 1882 : 318 ; Krebs, 1882 : 
295 ; Vosmaer, 1882: 20; 5. sp., idem, I.e.: 32, pi. i, figs. 22-23, pi. iv, figs. 140-144; 
5. domuncula, Marion, 1883: 65, 68; Carter, 1883: 30; Vosmaer, 1884: 121; idem, 
1885: 332; 5. montalbidus, Fristedt, 1885: 19, pi. iii, fig. 3; S. ficus, idem, I.e.: 20; 
5. virgultosa, idem, I.e.: 21; 5. suberia, Higgin, 1886: 86; S. lutkenii, Marenzeller, 
1886 : 3 ; 5. domuncula, Vosmaer, 1886 : 457 ; Thomson, 1887 : 241, pi. xvii ; Ridley and 
Dendy, 1887: p. xlv; 5. montalbidus, Fristedt, 1887: 428; Alcyonium compactum, 
Topsent, 1888: 134; Suberites domuncula, idem, I.e.: 134; 5. suberea, idem, I.e. 150; 
S. ficus, idem, I.e.: 150; idem, 1888: 1299; 5. domuncida, Lendenfeld, 1888: 65; 5. 
domunculus, Sollas, 1888 : 415 ; S. suberea, Hanitsch, 1889 : 158 ; S. ficus, idem, I.e. : 195 ; 
S. ficus, idem, I.e.: 195; Halichondria farinaria, Topsent, 1889: xxxviii; Suberites 
domuncula, Dendy, 1889: 56; Lendenfeld, 1889: 798; Topsent, 1890: 232; Hanitsch, 
1890: 195, 214; S. ficus, Hanitsch, 1890: 195, 216; 5. suberea, Topsent, 1890: 202; 
S. ficus, idem, I.e.: 202; S. farinaria, idem, I.e.: 203; S. ficus, Topsent, 1891: 14; 
idem, 1891: 127, 129; idem, 1891: 529; 5. domuncula, Hanitsch, 1891: 218; S. ficus, 



37o SUBERITES DOMUNCULA (OLIVI) 

idem, I.e.: 219; S.ficus, Topsent, 1892: 128; 5. latus, Lambe, 1893: 71, pi. iii, fig. 7, 
pi. v, fig. 7; 5. domunculus, Holt, 1892: 239; S.ficus, Levinsen, 1893: 410, fig. 21; 
S.farinaria, idem, I.e. : 412 ; fig. 22 ; 5. montalbidus, idem, I.e. : 413, fig. 23 ; 5. domun- 
cula, Celesia, 1893: 1, pis. v-viii; 5. virgultosus, Hanitsch, 1894: 177; 5. domuncula, 
idem, I.e. : 177 ; S.ficus, idem, I.e. : 177 ; S . farinarius , idem, I.e. : 177 ; S.ficus, Topsent, 
1894: 21, 23, 26; Halichondria f armaria, idem, I.e.: 21, 26; Suberites domuncula, 
idem, I.e.: 23; 5. suberea, Lambe, 1894: 126, pi. iv, fig. 3; 5. montalbidus, idem, I.e.: 
127, pi. iii, fig. 6 ; S.ficus, Weltner, 1894: 327 ; 5. heros, idem, I.e. : 328 ; 5. domuncula, 
Heider, 1895 : 283 ; 5. suberea, Lambe, 1895 : 126, pi. iv, fig. 3 ; 5. latus, idem, I.e. : 127 ; 
5. montalbidus, idem, I.e.: 127, pi. iii, fig. 6; S.ficus, Topsent, 1896: 275 ; idem, 1896: 
118; Lambe, 1896: 193, pi. ii, fig. 4; Ficulina ficus, Lendenfeld, 1897: 94, pis. iii, 
vi, vii, ix; Suberites domuncula, idem, I.e.: 118, pis. iv, vii, xi; Topsent, 1898: 126; 
Ficulina ficus , idem, I.e.: 129; Suberites domuncula, Thiele, 1898: 37; 5. heros, idem, 
I.e. : 37 ; 5. lutkenii, idem, I.e. : 38 ; 5. subereus, idem, I.e. : 38, pi. i, figs. 11-12, pi. viii, 
fig. 7 ; 5. placenta, idem, I.e. : 39, pi. viii, fig. 8 ; 5. sericeus, idem, I.e. : 39, pi. viii, fig. 
10 ; Prosuberites inconspicuus, idem, I.e. : 40, pi. viii, fig. 12 ; ? P. exiguus, idem, I.e. : 
40, pi. viii, fig. 13 ; Ficulina ficus, Topsent, 1899: 105 ; Ficulina ficus, idem, 1900: 203, 
pi. v, figs. 6-15; Suberites domuncula, idem, I.e.: 225, pi. vi, figs. 1-9; 5. suberea, 
Lambe, 1900: 161 ; S.ficus, idem, I.e. : 161 ; 5. montalbidus, idem, I.e. : 162 ; idem, 1900: 
24; idem, 1900: 277; 5. lutkenii, Topsent, 1900: 213; 5. domuncula, Cotte, 1901: 1; 
idem, 1901: 95; Bidder, 1902: 380; Ficulina ficus, Rousseau, 1902: 18, fig. 11; 
Suberella heros, Thiele, 1905 : 415 ; Suberites domuncula, idem, I.e. : 416 ; Swart- 
schewsky, 1905: 35, pi. ii, fig. 5, pi. iv, fig. 11, pi. vi, fig. 4; 5. heros, idem, I.e.: 
35; S. montalbidus, idem, 1906: 318, pi. xiii, fig. 3; Ficulina ficus, Lundbeck, 1907: 
559; idem, 1909: 453; Suberites sp., Arndt, 1912: 114; Ficulina ficus, Stephens, 
1912: 21; Massey, 1912 (see index p. 224 for page reference); Muller, 1913: 291; 
Topsent, 1913 : 25 ; F. lutkenii, idem, I.e. : 25 ; Stephens, 1915 : 35 ; Suberites domun- 
cula, Babic, 1921: 14; idem, 1922: 272; Ficulina ficus, Ferrer, 1922: 269; Suberites 
domuncula, Topsent, 1925: 633; Dembowska, 1926: 163; Ficulina ficus, Broch, 
1927: 5; Suberites domuncula, Topsent, 1928: 154; Ficulina ficus, idem, I.e.: 156; 
Arndt, 1928: 33, figs. 33, 34; Hentschel, 1929: 928; Suberites lutkenii, Hentschel, 
1929: 928; S. domuncula, Burton, 1932: 201 ; Vosmaer, 1933: 426, pi. i, figs. 2, 4, 12, 
16, pi. ii, figs, io-n, pi. iv, figs. 2, 10, pi. xviii, figs. 6-14, pi. xx, figs, n-13, pi. 
xxxvii, figs. 5-10 ; Burton, 1934: 313 ; Ficulina ficus, idem, I.e. : 313 ; Suberites lutkenii, 
idem, 1934 : 14 ; 5. domuncula, Topsent, 1934 : 14 ; Ficulina ficus, idem, I.e. : 16 ; Suberites 
domuncula, Arndt, 1935 : 39 ; Ficulina ficus, idem, I.e. : 39 ; Suberites domuncula, 
Burton, 1935: 77; C ho anites ficus, de Laubenfels, 1949: 19; Suberites domunculus, 
de Laubenfels, 1949: 20, figs. 16-18. 

Description of Species : Encrusting in young stages, later may assume one of two 
forms, either massive or globular, rarely lobate, and growing round an empty gastro- 
pod shell containing a hermit crab, or massive, globular, ficiform, clavate, or irregu- 
larly lobate ; surface even, finely hispid or harsh to touch ; texture firm ; oscules few, 
large, apical; colour, alive, white, greyish-white, white and red with blue patches, 
white marbled with blue and red, and various shades of yellow, orange, and red ; 



SUBERITES DOMUNCULA (OLIVI) 371 

skeleton a dense, irregular reticulation of tylostyli, 0-09 to 0-45 by 0-008 mm., with 
microstrongyla or microxea for microscleres, smooth or microspined, often sparingly 
present, 0-015 to 0-05 mm. long. 

Distribution: Throughout the Arctic Ocean, in the Atlantic Ocean north of o° 
latitude, and in the Pacific Ocean north of approximately 35 latitude. Bathymetric 
range from low-water springs to 1,331 m. (the optimum probably o and 90 m.). 

Ecology : Almost any kind of habitat, but more particularly on sandy or muddy 
bottom (presumably where gastropods or shells are likely to be present) . 



APPENDIX 
THE ECOLOGY OF SUBERITES DOMUNCULA 

Although Suberites domuncula, as now understood, has received so much attention 
in the literature, the data on bathymetric range and ecology are singularly meagre. 
This is true even where, as has happened several times, an author is reporting on a 
collection containing hundreds of specimens. There is, however, a series of observa- 
tions, given by Massy (1912), but as these are scattered over 215 pages and obscured 
by a wealth of faunistic data relating to other marine organisms, it has been thought 
worth while to abstract these and publish them in tabular form as an appendix. 

The identifications given in Massy (I.e.) were by Miss Jane Stephens, and one of the 
more interesting points to emerge is that in this series of trawlings off the coast of 
Ireland, comprising over 500 stations, sponges were obtained at more than 100 
stations, and the vast majority of these belonged to Suberites domuncula. Only a half- 
dozen other species were represented in the hauls, with a total of a dozen or more 
specimens. This substantiates the impression left by a study of the literature, as well 
as by personal experience, that the species is widespread over the continental shelf 
throughout its range and its population figures are comparatively high. It is, however, 
unfortunate that Massy should have been so indefinite on this last point. In describ- 
ing 'number of specimens taken' the words 'few', 'several', 'moderate', 'many' are 
far too indefinite. Had actual numbers been included, the list would have been so 
much more valuable. 

Summary of catches of Suberites domuncula recorded by Anne L. Massy off the coast 

of Ireland 







Number of 


Depth in 






Page 


Station 


specimens taken 


fathoms 


Nature of bottom 


Commensals 


3 


12 


1 


12-14 


sand and shells 


Eupagurus sp. 


15 


43 


few 


17-23 


fine sand 


E. cuanensis ? 


16 


44 


moderate 


25-27 


sand 


,, 


17 


45 


,, 


40-60 


,, 


— 


21 


57 


1 


48-60 


fine sand 


E. cuanensis 


26 


70 


several 


25-26 


fine sand and mud 


— 


28 


77 


2 


27-30 


sand and mud 


E. sp. 


29 


80 


few 


12-17 


mud and sand 


— 


3i 


83 


moderate 


I4f-i5* 


sand and shells 


— 


35 


102 


few 


12-16 


— 


E. sp. 



372 



SUBERITES DOMUNCULA (OLIVI 







Number of 


Depth in 






Page 


Station 


specimens taken 


fathoms 


Nature of bottom 


Commensals 


35 


104 


1 


14-16 


— 


E. sp. 


36 


107 


few (10 + ) 


20-23 


— 


E. sp. 


37 


108 


few 


13-14 


— 


E. sp. 


38 


113 


8 


21 


— 


E. sp. 


38 


114 


19 


21-25 


— 


2 with E. bernhardus; 17 on Denta- 
lium 


39 


116 


1 


16 


— 


E. sp. 


40 


118 


1 


21-23 


mud and sand 


E. sp. 


41 


122 


2 


11-13 


— 


E. sp. 


42 


126 


12 


43-60 


— 


10 with E. sp. ; 2 on Dentalium 


42 


125 


1 


12-14 


— 


E. sp. 


43 


129 


few 


13-15 


— 


E. sp. 


44 


131 


6 


21-28 


— 


Dentalium 


45 


135 


12 + 


9-10 


— 


,, 


46 


139 


2 


14-16 


— 


,, 


47 


143 


3 


17-20 


— 


E. sp. 


49 


146 bis 


1 


132-16 


— 


E. sp. 


53 


165 


1 


19-20 


sand and gravel 


E. sp. 


55 


173 


3 


13-16 


— 


E. sp. 


62 


198 


2 


48 


— 


E. sp. 


62 


199 


many 


18-24 


— 


E. bernhardus Aequipecten 


64 


203 


2 


— 


— 


E. bernhardus 


66 


206 


1 


11 


— 


— 


69 


216 


2 


12-19 


— 


E. sp. 


69 


217 


3 


32-50 


— 


E. sp. 


71 


222 


1 


15-16^ 


— 


E. cuanensis 


72 


224 


few 


44 


sand 


1 with E. cuanensis 


80 


248 


2 


10-12 


— 


i» »» 


83 


253 


3 


13 


— 


— 


85 


258 


1 


21-23 


mud 


— 


86 


261 


very scarce 


28 


fine sand and shells 


— 


87 


262 


1 


35-43 


sand 


— 


88 


264 


4 


17-23 


— 


— 


88 


265 


few 


242-25 


sand and shells 


E. bernhardus 


93 


280 


1 


8 


sand 


— 


96 


287 


6 


22 


fine sand and shells 


E. cuanensis 


96 


288 


9 


12I-132 


,, 


— 


97 


289 


2 


22-23 


mud and sand 


— 


97 


292 


2 


19-22 


sand and shells 


E. cuanensis ? 


104 


313 


1 


— 


— 


— 


106 


3i8 


1 


13 


coarse sand, gravel 


— 


107 


322 


moderate 


23 


sand 


E. cuanensis 


107 


323 


,, 


21^-23^ 


fine sand 


E. cuanensis and E. bernhardus 


109 


328 


2 


1 of 


fine sand, shells 


— 


112 


336 


3 


I4i-i7 


fine sand 


E. bernhardus 


119 


357 


1 


— 


— 


— 


124 


374 


1 


24-25 


sand 


— 


124 


375 


1 


232-24 


fine sand 


— 


136 


414 


2 


i6f-i9i 


,, 


— 


138 


418 


4 


23-232 


fine sand, shells 


E. cuanensis 


142 


438 


1 


8-8 £ 


,, 


— 


143 


439 


few 


192-232 


mud and sand 


E. bernhardus 


144 


443 


1 


13-19 


sand 


,, 


145 


444 


8 


222-24 


fine sand, shells 


E. sp. 


145 


445 


many 


25-26 


sand 


— 


146 


447 


2 


5-6 


,, 


— 


147 


45i 


2 


40-66 


mud, sand, shells 


E. bernhardus 



SUBERITES DOMUNCULA (OLIVI 



373 







Number of 


Depth in 






Page 


Station 


specimens taken 


fathoms 


Nature of bottom 


Commensals 


149 


455 


1 


14-15* 


fine sand, shells 


— 


153 


465 


1 


10 


fine sand 


— 


157 


476 


few 


23 


sand and shells 


E. sp. 


157 


477 


1 


24-25 


fine sand 


E. sp. 


161 


484 


1 


14-21* 


fine sand, shells 


E. bernhardus 


163 


487 


1 


19-23 


fine sand, mud 


— 


164 


491 


1 


7*"9 


fine sand 


— 


168 


500 


1 


IO-II* 


,, 


— 


168 


501 


1 


35-37 


mud 


— 


169 


504 


few 


42-46* 


mud and sand 


— 


171 


507 


1 


13-Mf 


fine sand, shells 


— 


173 


513 


1 


23^-25 


,, 


— 


173 


514 


several 


22-24 


sand 


E. bernhardus 


174 


515 


1 


22-26 


fine sand, shells 


— 


174 


5i6 


1 


19-22 


sand and shells 


— 


178 


526 


2 


7-7* 


sand 


— 


178 


527 


1 


10-13* 


,, 


— 


180 


532 


1 


14-Mf 


fine sand 


on shell 


181 


535 


2 


21-22* 


sand and shells 


E. sp. 


186 


545 


2 


i6*-i8£ 


mud 


— 


189 


553 


2 


41-52 


sand and shells 


E. bernhardus 


190 


554 


2 


14-19 


" 


— 



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Babic, K. 1921. Monactinellida und Tetractinellida der Adria. Glasn. hrv. prirodosl. 33: 77-93, 

9 ngs. 
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374 SUBERITES DOMUNCULA (OLIVI) 

Broch, H. 1927. Untersuchungen iiber die Marine-Bodenfauna bei Lindesness in Juni 1926. 

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1934. Sponges [in] Sci. Rep. Gr. Barrier Reef. Exped. 4: 513-621, 2 pis., 33 figs. 

1934- Observations on post-larval sponges of the genus Suberites. Ann. Mag. nat. Hist. 13: 

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1934. Zoological results of the Norwegian Scientific Expeditions to East Greenland. III. 

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1935. Some sponges from the Okhotsk Sea and the Sea of Japan. Explor. Mers russes, 22: 

61-79, 6 figs. 
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exp. gen. (3) 8: 1-33 1, 8 pis. 
19 1 3. Spongiaires provenant des campagnes scientifiques de la 'Princess Alice' dans les 

Mers du Nord. Result. Camp. sci. Monaco, 45: 1-67, 5 pis. 
1925. £tude de Spongiaires du Golfe de Naples. Arch. zool. exp. gen. 61: 623-725, 1 pi., 

27 rigs. 
1928. Spongiaires de l'Atlantique et de la M6diterranee. Result. Camp. sci. Monaco, 74: 

376 pp., 11 pis. 
1934. Sponges observers dans les parages de Monaco. (Premiere partie.) Bull. Inst. 

oceanogr. Monaco, 650: 1-42, 3 figs. 
Vosmaer, G. C. J. 1 88 1. Voorloopig berigt omtrent het onderzoek door den ondergeteekende 

aan de Nederlandsche werktafel in het Zoologisch Station te Napels verrigt. Nederl. 

Staatscourant, No. 109, 6 pp. 



378 



SUBERITES DOMUNCULA (OLIVI) 



Vosmaer, G. C. J. 1882. Report on the Sponges dredged up in the Arctic Sea by the ' Willem 

Barents'. Niederl. Arch. Z00L, Suppl. 1: 1-58, 4 pis. 
1884. Porifera [in] Bronn, Die Klassen und Ordnungen des Thierreichs, 2: 65-176, pis. iii, 

vii-xviii. 
1886. Porifera [in] Bronn, Die Klassen und Ordnungen des Thierreichs, 2: 369-496, pis. 

xxvi-xxxiv. 

1933- The Sponges of the Bay of Naples. Porifera Incalcarea. Capita zool. 5: 321-696, 

28 pis. 
Weltner, W. 1894. Spongien [in] Wiss. Meeresuntersuch. 1: 325-328. 
Wright, E. P. 1870. Notes on Sponges. Quart. J. micr. Sci. 10: 73-82, 3 pis. 



NOTES ON ASTEROIDS IN THE BRITISH 
MUSEUM (NATURAL HISTORY) 

III. 1 LUIDIA 

By AILSA M c GOWN CLARK 

(With Plates 39-46) 
The following species of the genus Luidia are represented in the Museum collection ; 
those of which the types are held are marked with an asterisk and those commented 
on in the text, with a dagger: 

aciculata Mortensen Hongispina Sladen 

*^ africana Sladen maculata Miiller & Troschel, with forma 

altemata (Say), with subspecies \numidica \herdmani forma n. 

Koehler magnified Fisher (f under aspera) 

*\aspera Sladen manritiensis Koehler 

atlantidea Madsen (f under africana) neozelanica Mortensen 

avicularia Fisher penangensis de Loriol 

bellonae Liitken phragma H. L. Clark 

ciliaris (Philippi) prionota Fisher 

clathrata (Say) \quinaria von Martens (incl. Himbata Sladen) 

*f Columbia (Gray) sarsi Duben & Koren (f under africana) 

elegans Perrier (f under africana) fsavignyi (Audouin) 

foliolata Grube *\ scotti Bell 

*\hardwickii (Gray) (incl. *forficifer Sladen) senegalensis (Lamarck) 

*heterozona Fisher tessellata Liitken (f under Columbia) 

Sladen's very full descriptions of the 'Challenger' material are excellent in them- 
selves, but examination of the type specimen of Petalaster hardwickii Gray shows 
that L. forficifer Sladen is a synonym of this. Gray's description was, as usual, very 
brief and inadequate in the light of the many species since described. His type 
specimen is accordingly dealt with in detail here, as are the types of Bell's species 
Luidia scotti from off Rio de Janiero. L. doello-juradoi Bernasconi (1941) seems to be 
identical with the latter. Sladen's types of Luidia aspera were found to include 
specimens of two other species, so that only the one described by him is left as the 
holotype. 

The very fine 'Siboga' report on Luidia by Doderlein (1920) provides a valuable 
subdivision of the genus and a comprehensive survey of the species known up to that 
time. The following species (see p. 380) have been described since 1920 or were not 
included by Doderlein. 

Doderlein's four main groups are most convenient for splitting up this unwieldy 
genus into more manageable units, but the limits between them are not absolutely 
sharp. For instance, L. scotti Bellbridges the gap between the Clathrata and Altemata 
groups. Also the subgenus Integr aster with such species as L. avicularia Fisher and 

1 Notes I and II appeared in Novit. Zool. 42 (1948) and Bull. Brit. Mus. (Nat. Hist.) Zool. 1 (4) 
(1950) respectively. 



3 8o 



ASTEROIDS IN THE BRITISH MUSEUM (NATURAL HISTORY) 



Name 


Locality 


Group 


moroisoana Goto, 1914: 301 


Japan 


Quinaria 


yesoensis Goto, 1914: 306 


,, 


,, 


superba A. H. Clark, 1917: 171 


Pacific coast of Colombia 


Alternata (?) 


porteri A. H. Clark, 1917a: 153 


Chile 


Ciliaris (?) 


scotti Bell, 191 7: 8 


Off southern Brazil 


Clathrata 


neo-zelanica Mortensen, 1925: 278 


New Zealand 


Ciliaris 


varia Mortensen, 1925: 275 


,, 


A Iternata 


aciculata Mortensen, 1933: 425 


St. Helena 


Ciliaris 


hexactis H. L. Clark, 1938 : 73 


NW. Australia 


Quinaria 


heterozona Fisher, 1940: 265 


W. Africa 


>> 


mortenseni Cadenat, 1941: 53 (= heterozona) 


,, 


,, 


doello-juradoi Bernasconi, 1941: 117 (= scotti) 


Argentina 


Clathrata 


patriae Bernasconi, 1941: 117 


,, 


it 


quequenensis Bernasconi, 1942: 253 


,, 


Alternata 


bernasconiae A. H. Clark, 1945: 19 (= alternata) 


NW. Atlantic 


,, 


atlantidea Madsen, 1950: 192 


W. and NW. Africa 


Ciliaris 



L. heterozona Fisher joins up the Quinaria and Ciliaris groups. Indeed, Fisher (1940: 
265) puts the last-named species actually in the Ciliaris group and Doderlein himself 
in his 'family tree' of the genus (p. 223) illustrates the link up of the two groups 
through the subgenus Integraster . 

As Mortensen (1925: 281) says, in discussing L. neozelanica, most of the species 
belonging to the Ciliaris group are distinguished by apparently trivial characters, 
coupled with their geographical location. This certainly applies to the three species 
L. sarsi from western Europe, L. atlantidea from West and North-West Africa, and 
L. africana from South Africa, in which the differing form and location of the pedicel- 
lariae and the size of the paxillar spinelets provide the main characters by which they 
can be recognized. 

However, it seems to me that too much importance has sometimes been placed on 
the occurrence or non-occurrence of pedicellariae as a specific character, rather than 
on their shape. For instance, Luidia sibogae Doderlein (p. 262) is based on a single 
juvenile specimen with R = only 19 mm., so it is not surprising that pedicellariae are 
only found in the interbrachial angles. The only other character in which it seems to 
differ from typical L. savignyi (Audouin) is in having unusually large spine-bearing 
paxillae, itself a somewhat variable feature in the latter species. Doderlein himself 
suggests that it may only be a young specimen of L. savignyi. Similarly I am doubtful 
of the specific value of L. mascarena Doderlein (1920 : 261) as distinct from L. savignyi 
also. The few specimens known from Mauritius and South-East Africa seem to have 
few, if any, ventro-lateral pedicellariae, but this is, in my opinion, at most a sub- 
specific distinction and anyway may not be borne out by a good series of adult 
specimens. 

At one time Luidia ciliaris (Philippi) was thought to have an Atlantic variety which 
was called normani, distinguished from the typical Mediterranean form by the pos- 
session of trivalved rather than bivalved ventral pedicellariae. However, even Lud- 
wig, the initiator of this variety, abandoned it on the evidence of further material, as, 
I suspect will also be the case with some of the other forms of Luidia. 

In the text that follows the reference lists quoted are not necessarily complete. 



ASTEROIDS IN THE BRITISH MUSEUM (NATURAL HISTORY) 381 

CLATHRATA GROUP 

Luidia Columbia (Gray) 

Text-figs. 1 and 2, Pl. 39, Fig. i 

Petalaster Columbia Gray, 1840: 183. 

non Luidia Columbia, H. L. Clark, 1910: 331, pl. 1, fig. 2; Doderlein, 1920: 253; Bernasconi, 1943: 7, 

pl. 4, figs. 2 and 3 (= L. tessellata Liitken). 
Luidia brevispina Liitken, 1871: 288; Doderlein, 1920: 253, figs. 10, 14, and 22. 

Type: R/r = 58-65 mm./i2 mm. = 5/1. San Bias. Cuming collection. 

The specimen is dry and not in a very good condition. The ventral side seems to 
have been coated in glue particularly at the interbrachial angles which are distorted. 
Most of the spines, short as they are, have become adpressed to the surface or 
broken off. 

Note : Gray has obviously assumed that the specimen came from the San Bias on 
the Atlantic side of the isthmus of Panama, which was at that time part of Colombia, 
hence his specific name. I am unable to trace any place called San Bias on the Pacific 
coast of Colombia, but there is a town of that name on the west coast of Mexico near 
Mazatlan, where other similar specimens have been taken (the types of L. brevispina 
Liitken) . Since Cuming only collected on the west coast of Central America and some 
shells from his collection are recorded as coming from 'San Bias, Gulf of California' 
it is presumably from there that this specimen came. 

Diagnosis. A species of Luidia belonging to the Clathrata group of Doderlein, with 
two rows of lateral paxillae forming transverse rows with the larger supero-marginal 
series ; dorsal paxillae with large, flat, polygonal, central granules surrounded by much 
more slender peripheral spinelets ; no pedicellariae ; one very short, tapering marginal 
spine just below the ambitus on each infero-marginal plate, with two shorter flattened 
ones above it ; ventral infero-marginal spines very short and squamiform ; the single 
ventro-lateral plates hardly projecting from underneath the inner ends of the corre- 
sponding infero-marginals ; three relatively short, thick adambulacral spines. 

Description. The largest dorsal paxillae are the supero-marginals which are 
proximally wider than long but distally become square. They form transverse series 
with the two outermost lateral rows of paxillae, which are square (or slightly wider 
than long) proximally, becoming relatively longer distally. Towards the middle of the 
rays the paxillae become smaller and more irregularly arranged, having about seven 
flat, polygonal, central granules as compared with the twelve or so of the supero- 
marginal series. The peripheral spinelets around the paxillae are much more slender. 

There are no pedicellariae on either side. 

The infero-marginal plates are, as usual, very short and raised into a ridge extend- 
ing a little way on to the dorsal side, where they bear a few short stumpy spinelets. 
On the ambitus, or just above it, are two (rarely one), short, flattened spines, ex- 
panded outwardly rather like a hoof seen in side view. Below these comes a single 
short tapering spine, about half as long again as the two above it but, even so, not as 
much as 1 mm. in length. On the ventral side there are two somewhat irregular rows 
of expanded, squamiform spinelets, with smaller peripheral ones on either side. 



382 ASTEROIDS IN THE BRITISH MUSEUM (NATURAL HISTORY) 

The ventro-lateral plates are largely overlain by the infero-marginals so that only 
a small lobe protrudes. It is impossible to tell how many there are in the inter- 
brachial angle owing to the poor condition of the specimen. They do not appear to 
bear any distinct armature, although possibly they may carry a squamiform spinelet 
similar to and consecutive with the infero-marginal spinelets. 

The adambulacral plates have the usual curved, compressed furrow spine followed 
by two other spines, the middle one being slightly curved at the base, otherwise 





2 m 

Fig. i. 



Fig. 2. 



Text-fig. i. Luidia Columbia (Gray). Type. Dorsal view of two infero-marginal plates and the 
adjacent paxillae from the proximal part of an arm. 

Text-fig. 2. Luidia Columbia (Gray). Type. Ventral view of one side of two segments, that on 
the left having been treated with sodium hypochlorite. (The arrow points towards the mouth.) 



cylindrical and gently tapering, while the outer one is stouter and a little shorter. 
There may be several spinelets along the adoral edge of each adambulacral plate, of 
which one on a level with the outermost large spine may be enlarged occasionally. 

There is a faint tinge of greenish colour on the dorsal side. 

Remarks. Liitken, H. L. Clark, and Doderlein have all had a mistaken impression 
of this species, which is hardly surprising after Gray's very brief diagnosis, for Liitken 
when describing L. tessellata (1859: 40) from Puntarenas (on the west coast of Costa 
Rica) queried it as a possible synonym of Luidia Columbia, which it is not, and later 
(1871: 288) described as a separate species L. brevispina from Mazatlan, Mexico, 
which is clearly identical with L. Columbia. 



ASTEROIDS IN THE BRITISH MUSEUM (NATURAL HISTORY) 383 

There are two specimens in the British Museum identified as Luidia tessellata by 
Liitken and labelled as coming from Realejo, Punt arenas (the type locality). These 
fully agree with the description Doderlein has given for Luidia Columbia (p. 253), 
having long slender marginal and adambulacral spines. The longer of the two ambital 
spines (the lower one) is 3-5-4 mm. in length, while the upper one is usually about 
2 mm. long. The adambulacral spines are about 3 mm. long and with the slender 
spines on the ventral surface of the infero-marginal plates give the under side a 
'shaggy' appearance quite distinct from that of Luidia Columbia with its very 
abbreviated armature. 

Luidia tessellata is then a valid species and it is L. brevispina which is the synonym 
of Luidia Columbia (Gray) . 

As for Luidia marginata Koehler (1911a: 17) from Mazatlan, Doderlein (p. 251) says 
that it differs from L. brevispina (i.e. Columbia) in having numerous interradial ventro- 
lateral plates in the interbrachial angles, although Koehler himself makes no mention 
of this. It is unfortunate that the type of L. Columbia is in such a condition in this 
region that no comparison can be made. 

ALTERNATA GROUP 

Luidia scotti Bell 

Text-fig. 3 ; Pl. 40, Fig. i 

Luidia scotti Bell, 191 7: 8. 

Luidia doello-juradoi Bernasconi, 1941: 117; 1943: 8, pl. 1, fig. 3, pl. 2, figs. 2-3, pl. 3, figs. 4-5. 

St. 42. 'Terra Nova' Expedition. 22 56' S. : 41 34' W. (off Rio de Janeiro). 73 m. 
15 specimens. 

Holotype selected by A. M. Clark with R = 60 mm., r = 8 mm., R/r = 7-5/1, 
br. = 9 mm., British Museum registered number 1915.2.1.64. 

Diagnosis. A species of Luidia linking the Alternata and Clathrata groups, with two 
lateral rows of paxillae forming transverse rows with the supero-marginal series ; no 
dorsal pedicellariae but three- or four-valved ones are present on most of the ventro- 
lateral plates in the interbrachial angles and at the bases of the arms ; one large 
marginal spine at the ambitus with a smaller one above it and four or five others 
below on the ventral face of the plate, all of them much smaller than the ambital 
spine ; four adambulacral spines, the outermost two placed on a line parallel to the 
furrow. 

Description. The sides of the rays are almost vertical up to the second row of 
paxillae in from the supero-marginal series. The centre of the disk and rays is quite 
flat. The madreporite is concealed. 

The supero-marginal paxillae are square or slightly longer than wide. Forming 
transverse rows with them are two series of lateral paxillae, of which the outer row, 
at least, are wider than the supero-marginals. Across the middle of the ray there are 
about thirteen rows of less regularly arranged plates, which become progressively 
smaller towards the mid-radial line. The small central plates, both of the disk and the 
arms, bear three or four short, thick, spaced paxillar spinelets surrounded by about 
twelve thinner peripheral ones. The number of spinelets on each paxilla increases 

zoo. 1, 12. 2 E 



384 



ASTEROIDS IN THE BRITISH MUSEUM (NATURAL HISTORY) 



towards the sides of the rays to the outermost lateral series, each plate of which bears 
about twelve central and thirty peripheral spinelets. All the paxillar spinelets have 
the rounded tops distinctly thorny under magnification. 
There are no dorsal pedicellariae. 

The infero-marginal plates are mainly ventral in position but have a small area 
covered with paxillar spinelets on their uppermost edge at the side of the arm. Below 
this, at the ambitus of the ray, is a large, pointed, curved spine about 2-5 mm. in 

length. Above this is a smaller spine, usually about 1 mm. 
long, although rarely it is two-thirds as long as the 
ambit al spine. On the ventral side of each infero-mar- 
ginal plate is a series of four or five much smaller 
pointed spines, slightly flattened, one being occasionally 
replaced by a pedicellaria as in the figure. The outermost 
is the largest and measures just over 1 mm. in length. On 
each side of this row there may be a series of smaller, 
stumpy spines, while on the edges of the plates are the 
usual fringing spinelets. 

The adambulacral plates have a curved, sabre-like 
furrow spine backed by a larger tapering one, followed in 
turn by a pair of spines of which either the adoral may 
be smaller while the other is the same size as the second 
spine, or else both of them are smaller. On the outer 
edge of the plate lie one or two smaller spines or 
spinelets. In the interbrachial angle each ventro-lateral 
plate bears a three- or f our-valved pedicellaria, but these 
only extend on to the proximal part of the ray up 
to about the sixth joint, beyond which there are only 
spinelets. There are no pedicellariae on the mouth plates. 
The colour has been lost in the type after thirty-five 
years in spirit, but some other specimens are dark brown 
along the middle of the rays while a small one has 
brown blotches at intervals across the arms, as Bell 
described when the material was fresh. 

Variability. A paratype slightly larger than the specimen described above has the 
ventro-lateral pedicellariae extending on to most of the plates in the proximal half of 
the arm, not just on those in the interbrachial angle. 

The long narrow arms with an R/r ratio of 7 (or more)/i are found in all the speci- 
mens from this station. 

Remarks. Bell compared this species with Luidia clathrata (Say), but the rela- 
tively smaller supero-marginal paxillae compared to the outermost lateral series and 
the presence of pedicellariae and blotched coloration readily distinguish his own 
species. These characters are more typical of the Alternata group of Doderlein than 
of the Clathrata group. 

The affinities of Luidia scotti are obviously with the species included in Doderlein's 
subgenus Armaster, particularly L. armata Ludwig (1905: 85) and L. ludwigi Fisher 




Text-fig. 3. Luidia scotti Bell. 
Type. Ventral view of one side 
of two arm segments, that on 
the left having been treated 
with sodium hypochlorite. 
( The arrow points towards the 
mouth.) 



ASTEROIDS IN THE BRITISH MUSEUM (NATURAL HISTORY) 



385 



(1906a and 1911). However, without Pacific material for comparison it cannot be 
decided whether the forms on both sides of South and Central America represent the 
same species. They are certainly very closely related. 

Luidia scotti is obviously identical with L. doello-juradoi Bernasconi (1941) from the 
mouth of the river Plate. The type of that species also has supero-marginal paxillae 
equal in length to, but not so wide as, the outermost lateral row and three- or four- 
valved pedicellariae on the ventro-lateral plates. The only difference appears to be that 
the two marginal spines are almost equal in length in L. doello-juradoi, whereas in the 
types of L. scotti the upper one is usually less than half the size of the lower. 

It is unfortunate that Bell's description was so brief and omitted any mention of 
the distinctive pedicellariae. 



Luidia savignyi (Audouin) 
Pl. 40, Fig. 2 

Asterias savignyi Audouin, 1826. 

Luidia savignyi, Gray, 1840: 183 ; Perrier, 1875: 342 ; Koehler, 1910: 10, pl. 1, fig. 5, pl. 6, fig. 3. 

Luidia mascarena Doderlein, 1920: 261, fig. 5. 



Number 


B.M. Reg. No. 


Locality 


Source 


1 
1 
2 

1 
1 


1903.4.2.39 
1904.3.3.66 
69.6.25.9-10 
1951.1.6.1 
1927. 1. 10.90 


Wasin, N. of Zanzibar, 10 fms. 

Pearl Bank, Ceylon 

Gulf of Suez 

28 32' S. : 32 26' E., NE. of Durban, 20 m. 

Suez 


Crossland collection 

Herdman collection 

R. McAndrew 

Cape Town University 

C.U. Suez Canal Expedition 



Remarks. Luidia mascarena Doderlein, from Mauritius, apparently only differs from 
L. savignyi in lacking ventro-lateral pedicellariae. This is the case with Doderlein's 
two specimens and with de Loriors one, according to Koehler. The specimen from 
north-east of Durban, with R = 140 mm., has a single pedicellaria on each side of 
each interbrachial angle, usually on the third or fourth plate, and three or four others 
farther out on the arm. The Wasin specimen is unfortunately juvenile with R = only 
35 mm. It has a total of three ventral pedicellariae, of which two are in one inter- 
brachial angle. The Suez specimen collected by the Cambridge University Expedition 
(R = 38 mm.) has no ventral pedicellariae at all, as remarked on by Mortensen 
(1926: 121), although the other two from the same locality, collected by McAndrew, 
both have pedicellariae on most of the ventro-lateral plates. They are, however, 
much larger (R === 95 mm. or more) . 

The specimen from Pearl Bank, Gulf of Manaar (Pl. 40), is that 'with spines on the 
surface of its rays ', this comment of Bell's being reproduced under Luidia hardwicki in 
Herdman's report (1904: 143). The dorsal spines certainly are numerous and very 
large, measuring about 4 mm. in length. R = 50 mm. Coupled with the powerful 
dorsal armature, the ventral spines and pedicellariae are unusually long for the 
species. But for the degree of development of the spines there seems to be little 
difference between this form and typical L. savignyi. More material is needed to 
show whether it comes within the range of variation of the latter. 

The dark patches on the arms of this species when seen through a lens are shown to 



386 ASTEROIDS IN THE BRITISH MUSEUM (NATURAL HISTORY) 

be produced by pigmentation on the surfaces of those paxillae which come within the 
tinted area, extending on to the bases of the paxillar spinelets. This throws into sharp 
contrast the white tips of the spinelets. In the smaller specimens there is often only 
a single central spinelet on the mid-radial paxillae. 

All of these specimens are seven-rayed in marked contrast to the two five-rayed 
ones from Madagascar remarked on by Koehler (1910: 14) in his own collection. He 
says 'les pedicellaires sont particulierement abondants', which does not seem to be 
the case in the few seven-rayed specimens of L. savignyi known from Mauritius on 
one side and the coast of South-East Africa on the other. Koehler could not find any 
other character by which to separate this five-rayed form from the more widespread 
seven-rayed one. 

The very large nine-rayed specimen from Mauritius recorded by Bell (1889: 422) 
and purchased from M. de Robillard is not L. savignyi but L. mauritiensis Koehler 
(1910: 15, pi. 1, figs. 6-7), a species more nearly related to L. magnified Fisher, 
from the Hawaiian Islands with ten arms and L. aspera Sladen from the Philippines, 
with eight, also having dorsal spines on many consecutive plates. A second specimen 
actually had ten rays originally, but all have been broken off and nine pieces splinted 
on to the disk neglecting to leave a gap, so that from the dorsal side nine appears to 
be the actual number. It is dried and altogether in a bad state. 

Luidia aspera Sladen 
Text-figs. 4 and 6 
Luidia aspera Sladen, 1889: 248, pi. 43, figs. 1-2, pi. 45, figs. 9-10. 

Of the original four types of this species, the two young ten-rayed specimens, each 
with R = about 40 mm., and a slightly longer odd arm, are obviously not the same 
as the two large ones from Zamboangan, in the Philippines, in 10 fathoms, as for one 
thing they do not have blotched coloration. These two small specimens, from 
'Challenger' stations 204, off Tablas Island, Philippines, in 100 fathoms, and 209, 
north of the Admiralty Islands in 150 fathoms, are Luidia avicularia Fisher, a 
species belonging to the Quinaria group. 

The remaining two specimens, one with eight, the other with ten rays, are otherwise 
superficially similar, having blotched coloration and several rows of lateral paxillae 
with erect spines on many consecutive plates. However, closer examination shows 
that they differ in a number of ways. 

Both of them have R = c. 170 mm., but in the ten-rayed specimen the disk dia- 
meter is 45 mm. and in the eight-rayed one only 35 mm. The latter has much longer 
and more slender three-bladed pedicellariae, numbering at the most two to each seg- 
ment, whereas in the ten-armed one there are three ventrolateral plates and corre- 
spondingly three pedicellariae (at least proximally), which are also shorter and more 
abruptly tapering. This is obviously a specimen of Luidia magnifica Fisher (1906: 
1033), the type of which, also ten-rayed, came from the Hawaiian Islands in 43-73 
fathoms. The eight-rayed specimen, which Sladen described and figured, is thus left 
as the only type of Luidia aspera. 





Fig. 4. Fig. 5. 

Text-fig. 4. Luidia aspera Sladen. Type. Dorsal view of the upper ends of two infero-marginal 
plates and the adjacent paxillae. 

Text-fig. 5. Luidia magnified Fisher. Dorsal view of the upper ends of two infero-marginal 
plates with the adjacent paxillae, showing the papulae between the plates. 




Fig. 6. Fig. 7. 

Text-fig. 6. Luidia aspera Sladen. Type. Ventral view of one side of two segments, that on the 
left having been treated with sodium hypochlorite. (The arrow points towards the mouth.) 
Text-fig. 7. Luidia magnifica Fisher. Ventral view of one side of two segments, that on the left 
having been treated with sodium hypochlorite. (The arrow points towards the mouth.) 



388 ASTEROIDS IN THE BRITISH MUSEUM (NATURAL HISTORY) 

The differences between the two species can be listed as follows : 

L. aspera L. magnified 

1. Eight rays. i. Ten rays. 

2. Dorsal spines present on the paxillae of the 2. Dorsal spines present on the paxillae of the 
third to the fifth (sixth) lateral rows, count- second to fourth (fifth) lateral rows, count- 
ing in from the supero-marginal series. ing in from the supero-marginal series. 

3. No pedicellariae on the supero-marginal 3. Pedicellariae present on the supero-mar- 
plates. ginal and some of the first lateral series of 

paxillae. 

4. Ventral pedicellariae about three times as 4. Ventral pedicellariae only twice as long as 
long as their basal width. their basal width. 

5. Two ventro-lateral plates occur on each side 5. Three ventro-lateral plates present on each 
of each segment. side of each segment. 

6. Three large adambulacral spines. 6. Two large adambulacral spines. 

7. Furrow spine long. 7. Furrow spine rather short. 

The accompanying comparative illustrations (Text-figs. 4-7) of these two speci- 
mens help to emphasize these differences. 

The occurrence of these two species together suggests that the eight-rayed Luidia 
hystrix Fisher (1906: 1032), also from the Hawaiian Islands in depths of 14-52 
fathoms, is probably identical with L. aspera. The differences mentioned by Fisher 
are that in L. aspera only three rows of lateral paxillae are spiniferous and there are 
only three adambulacral spines but two pedicellariae on many segments, whereas in 
L. hystrix nearly all the dorsal paxillae are spiniferous, there are four adambulacral 
spines, and pedicellariae only occur on about half the segments and then never more 
than one at a time. I believe these three differences are all subject to variation, but 
to what extent can only be settled by further material. 

The minor differences between the 'Challenger' specimen of L. magnifica and the 
type of that species, which has R = 330 mm., are all in my opinion attributable to 
the great size of the latter. 

The seven-rayed specimen from Mozambique, recorded as Luidia aspera by Simp- 
son and Brown (1910 : 49) clearly belongs to L. savignyi (Audouin), as noted by Fisher 
(1919: 171). 

Luidia aspera is certainly very closely related toL. savignyi, and apart from having 
eight rays rather than seven, the only notable difference seems to be that L. aspera 
has relatively small dorsal spines occurring on many consecutive lateral paxillae, 
whereas in L. savignyi the spines are rather more robust and usually only occur 
sporadically on the lateral series of paxillae. 

Luidia alternata numidica Koehler 
Pl. 41, Fig. i 

Luidia numidica Koehler, 1911: 3, pl. 1, figs. 8-1 1. 

Luidia alternata var. numidica Madsen, 1950: 206, text-fig. 9. 

There are five specimens of this subspecies in the Museum collections, of which 
one from Boa Vista Island in the Cape Verde group, collected by Crossland, has particu- 
larly numerous spine-bearing paxillae in the second and third (rarely in the fourth) 



ASTEROIDS IN THE BRITISH MUSEUM (NATURAL HISTORY) 389 

rows inwards from the supero-marginals. On the other paxillae the peripheral 
spinelets are distinctly more slender than the shorter central ones, as in typical West 
Atlantic L. alternata, not like the type of numidica. Indeed, this specimen is very 
near typical Luidia alternata. 



Luidia maculata forma herdmani forma n. 
Text-fig. 8 ; Pl. 41, Figs. 2 and 3 

Pearl Bank, Gulf of Manaar, Ceylon. Herdman collection. 1904.3.3.8-9. 3 speci- 
mens (2 very young) . 

Tuticorin, Madras. Thurston collection. 88.1.2.64. J specimen. 

Type: larger specimen from Pearl Bank. R/r = 46 mm./8'5 mm. = 5*4/1. 

One arm has been broken and has partly regenerated. 

Diagnosis. A form of Luidia maculata differing from the typical form in having 
only six arms and the paxillae of the disk and proximal parts of the rays with a knob- 
like enlarged central spinelet about twice the height of the peripheral ones. 

Description. The dorsal side is convex and only the 
centre of the disk is at all flattened. The madreporite is not 
visible. The dorsal paxillae of the outermost lateral row are 
the largest, being even larger than the supero-marginal 
series. Whereas the latter are square, the lateral paxillae 
are wider than long. Proximally four lateral series also form 
regular transverse rows with the supero-marginals, but 
midradially there are two rows of more numerous, smaller 
plates. Four to seven somewhat polygonal, thick, granule- 
like central spinelets and about 14 thinner peripheral spine- 
lets cover each small paxilla of the centre of the arm but on 
the outermost lateral series there are about 12 central and 28 
peripheral spinelets. On the disk also the paxillae are smaller 
than those on the sides of the rays as well as being round 
rather than rectangular. They have 1-3 (most commonly 1) 
very enlarged, round-topped central spinelets surrounded by 
about 6 less robust ones with the still more slender peripheral 

spinelets alternating in position with these. The central spinelet usually projects 
somewhat above the top of the other spinelets. Such paxillae only occur on the 
disk and at the bases of the arms and their round-topped central spinelets 
are never comparable to the paxillar spines of such species as Luidia savignyi 
( Audouin) . 

The infero-marginal plates lie almost entirely on the ventral side. Their longest 
spine is near the upper edge of the plate and reaches a maximum of 2-5 mm. in 
length. Proximally there may be a smaller spine dorsal to this one, contributing to 
the fringe of spines projecting laterally from the ambitus of the ray. On the ventral 
side are three, rarely four, equal-sized, pointed, erect spines, considerably shorter than 
the upper large one. A row of smaller spinelets runs down each side of the plate with 




Text-fig. 8. Luidia 
maculata var. herdmani 
var. n. (a) Paxilla of the 
outermost lateral series, 
(b) lateral, and (c) dorsal 
views of a disk paxilla. 



39Q ASTEROIDS IN THE BRITISH MUSEUM (NATURAL HISTORY) 

a few scattered ones between the larger spines. As on the dorsal paxillae there are no 
pedicellariae on the infero-marginal plates. 

The adambulacral plates have the usual curved, compressed, sabre-like furrow 
spine, followed by a larger spine also compressed laterally but which appears to be 
widened near the tip when viewed from a direction parallel to the furrow. A third 
slightly shorter and narrower spine stands behind this one backed by a three- or even 
four-valved pedicellaria on the outer edge of the plate. Adoral to the biggest spine 
there may be a single spinelet like those bordering the infero-marginal plates. 

There are no pedicellariae in the furrow on the mouth plates. 

The coloration, like that of typical L. maculata Muller & Troschel, is blotched. In 
spirit there are pairs of dark brown marks along the rays. Another specimen has a 
six-pointed brown star on the centre of the disk and the two small ones have a star 
effect on the disk caused by a V-shaped dark mark in each interbrachial angle. Many 
of the ventral spines and spinelets are dark-tipped. 

Variations. In the Tuticorin specimen the pedicellaria outside the adambulacral 
spines is more often than not absent. Also the enlarged central spinelet on the paxillae 
at the base of the arms may be as much as three times as long as the other spinelets. 

The small specimens (R = c. 15 mm.) have very thorny-tipped peripheral spine- 
lets on the dorsal paxillae. 

Remarks. This form only seems to differ from typical L. maculata in having 6 arms 
rather than 7-9, besides the conspicuous enlargement of the central granule of the 
disk paxillae. The latter feature was not encountered by Doderlein and does not 
occur in the British Museum material of L. maculata, although Fisher (1919 : 169) says 
that in eight-armed Philippine specimens the central spinelet is often larger than the 
others which become progressively smaller towards the periphery of the paxilla. The 
two forms seem to overlap in their ranges as typical specimens of L. maculata, with 
seven or eight arms and uniform central paxillar granules, have been taken at 
Tuticorin and on the Pearl Bank, off Ceylon. Unfortunately, no details of locality 
were recorded at the time. Luidia maculata usually has pedicellariae on the marginal 
and dorso-lateral paxillae but Doderlein says that their presence is very variable and 
they may be completely absent, as here. 

Koehler (1910a, pi. 15, fig. 2) shows a figure of the ventral side of a six-armed speci- 
men of L. maculata from the Moluccas, but that number seems to have been rarely 
recorded. 

The consistent combination of the two features — presence of only six arms and 
enlargement of the central spinelet of the disk paxillae — seems to be sufficient grounds 
for giving this form a special name. 

From Luidia penangensis de Loriol, a six-armed species also from the Indian Ocean, 
with an enlarged spinelet in the middle of each paxilla (although not just on the disk 
and arm-bases), this form can be told at a glance by the absence of a conspicuous 
madreporite as well as by all the other characters — such as the occurrence of two-bladed 
pedicellariae on the mouth plates — which distinguish the Quinaria group (to which 
L. penangensis belongs), from the Alternata group. 



ASTEROIDS IN THE BRITISH MUSEUM (NATURAL HISTORY) 391 

QUINARIA GROUP 

Luidia hardwicki (Gray) 

Pl. 39, Figs. 2 and 3 

Petalaster hardwickii Gray, 1840: 183. 
Luidia hardwickii, Perrier, 1875: 331 (1876: 251). 

Luidia forficifer Sladen, 1889: 258, pl. 44, figs. 5 and 6, pl. 45, figs. 5 and 6; Doderlein, 1920: 
278, text-fig. 3, pl. 20, figs. 28 and 29. 

Type: R = 32 mm., r = 5-5 mm., R/r = 6/1, br. = 5-5 mm. Registered number 
1938.5. 12. 12. Indian Ocean. 

Diagnosis. A species of Luidia belonging to the Quinaria group, with two or three 
lateral series of paxillae forming transverse rows with the supero-marginals ; large 
pedicellariae present on the mouth plates and on the outer part of the adambulacral 
plates ; a single enlarged marginal spine at the top of each infero-marginal plate, with 
smaller appressed spines on the ventral side of the plate. 

Description. Three, distally two, rows of lateral paxillae form transverse rows 
with the supero-marginal series. The inner paxillae are progressively smaller towards 
the mid-radial line. At the base of the arm there are fifteen paxillae across the width, 
including the two supero-marginal series. Those in the middle of the ray are, of 
course, more numerous than the lateral ones, but also tend to be arranged in trans- 
verse and longitudinal rows. 

These smaller paxillae, both in the centre of the disk and along the rays, have 
1-5 central spinelets, resembling slightly elongated granules. On the arms the number 
is more commonly one and this one may be a little enlarged. On each small paxilla 
there are also 10-12 peripheral spinelets, 2-3 times as long as wide and only slightly, 
if at all, thinner than the central ones. The supero-marginal paxillae have up to 
10 short central spinelets and about 20 longer peripheral ones. The outermost lateral 
series have about 8 central and 16 peripheral spinelets. 

Pedicellariae seem to be absent from the dorsal side. 

The madreporite is concealed by the paxillae. 

The ventral side has suffered somewhat in drying, but there is a pair of very long 
pedicellariae projecting over the furrow from each mouth angle, about 1-5 mm. in 
length and very similar in size to the larger adambulacral spines. There is a curved, 
sabre-like furrow spine on each adambulacral plate backed by the usual longer, 
stouter spine and another spine not as large as the second ; the three form a straight 
row at right angles to the furrow. Adoral to the two outer spines is a very big pedi- 
cellaria, about two-thirds as long as the longer spine and with the blades of the valves 
almost as broad at the tip as at the base (like those of L. forficifer as figured by 
Doderlein (1920: 278, text-fig. 30)). 

The ventro-lateral plates are very small, each one forming a little semicircle at the 
inner end of an infero-marginal plate. They are either bare or only have a few small 
spinelets. The infero-marginals have two or three appressed spines in a series down the 
middle of each plate, with smaller spinelets on each side. At the ambitus, which comes 
just below the small cluster of paxillar spinelets at the uppermost edge of the plate, 
is a single spine measuring about 1-5 mm. in length and just under 0-5 mm. in width 

zoo. 1, 12. 2 F 



392 ASTEROIDS IN THE BRITISH MUSEUM (NATURAL HISTORY) 

at the base. Some pedicellariae, about half the size of those on the adambulacral 
plates, occur on the infero-marginals. 

Remarks. This description agrees very closely with Sladen's of L. forficifer (1889 : 
258) with the exception of the infero-marginal plates, which, in the latter, have five 
squamiform spinelets in a row, whereas in the type of L. hardwicki, these spinelets 
are fewer and less regularly arranged. The smaller size and poor condition may 
account, at least partly, for this. 

Although the type of L. forficifer irora 'Challenger' station 187 (Booby Island, Torres 
Strait) has no pedicellariae on the infero-marginal plates, some are present in a 
larger co-type from station 188 (Arafura Sea near Torres Strait). There is then no 
character differing to an extent sufficient to separate the two specifically, so that 
forficifer becomes a synonym of Luidia hardwicki (Gray) . 

British Museum specimens named L. hardwicki by Bell, from Macclesfield Bank in 
the South China Sea, have pedicellariae on many of the dorsal paxillae, not just on the 
proximal marginal ones as in L. quinaria. Such pedicellariae are absent in the 
types of both L. hardwicki and L. forficifer, but their presence in other species of 
Luidia is very variable and their occurrence cannot be used as a specific character. 
These Macclesfield Bank specimens also have relatively few adambulacral plates 
bearing pedicellariae. 

Two specimens from the Great Barrier Reef Expedition, named by Livingstone 
L. forficifer, also have some pedicellariae on the dorsal paxillae. 

Luidia quinaria von Martens 
Text-fig. 9 

Luidia maculata var. quinaria von Martens, 1865: 352. 

Luidia quinaria, Ives, 1891: 211 pi. 9, figs. 5-9; Doderlein, 1920: 244, 275, text-fig. I. 

Luidia limbata Sladen, 1889: 251, pi. 44, figs. 3-4, pi. 45, figs. 7-8. 

One of the ten specimens of L. quinaria in the British Museum comes from Hako- 
date, in northern Japan. The pedicellariae on its mouth plates 
are rather thick, approximating in shape to those of L. amu- 
rensis Doderlein (1920: 277, text-fig. 2), from Vladivostok, 
which is in almost the same latitude as Hakodate. . 

Also in this specimen, as in L. amurensis, the pedicellariae 
on the marginal paxillae are little bigger than the central 
granules, not conspicuously larger as in specimens of L. 

I I quinaria from southern Japan. 

■mm. The three types of Luidia amurensis completely lack pedi- 

Text-fig. 9. Luidia cellariae on the adambulacral and ventro-lateral plates, but 
quinaria von Martens. th are present on most of the adambulacral plates in this 
Pedicellariae from the TT : , , , . ... . ,. .. .. . , 

mouth plates of a sped- Hakoda te specimen, although these pedicellariae are also 

men from Hakodate, relatively thicker than those of L. quinaria figured by Doder- 
northern Japan. lein (p. 272, text-fig. lb). 

The two forms are obviously very closely related, and 
L. amurensis may be better placed as a northern subspecies of L. quinaria. 




ASTEROIDS IN THE BRITISH MUSEUM (NATURAL HISTORY) 



393 




CILIARIS GROUP 

Luidia africana Sladen 

Luidia africana Sladen, 1889 : 256, pi. 44, figs. 1 and 2, pi. 45, figs. 1 and 2 ; Mortensen, 1933a : 239, 
text-figs. 3 and 4; Madsen, 1950: 188, text-fig. 4, pi. 16, figs. 3 and 4. 

The types of this species are four specimens from Simon's Bay, South Africa, and 
one from the coast of Morocco, near Gibraltar. The latter is broken into separate 
arms and the only complete specimen is a 

South African one with R = 160 mm. This is ■ — ■ 

the one figured in Sladen's plate 44, although 
Madsen suspected those illustrations were of 
the Moroccan specimen, which he thought was 
more likely to be Luidia atlantidea Madsen 
(1950: 192). Neither of these suppositions is 
correct. The light stripe along the sides of the 
arms in the figure of the dorsal side is probably 
an illusion created by the comparison with the 
darker mid-radial line. It is impossible to tell 
after so long in spirit whether such a white 
stripe, like that of L. atlantidea, ever existed 
in